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Review

Distributional Range Shifts Caused by Glacial–Interglacial Cycles: A Review on Timing, Main Processes, and Patterns of Late Pleistocene Marine Dispersal by Invertebrates in the NE Atlantic

by
Sérgio P. Ávila
1,2,3,4,5
1
MPB—Marine Palaeontology and Biogeography Lab, University of the Azores, Rua da Mãe de Deus, 9501-801 Ponta Delgada, Açores, Portugal
2
CIBIO—Açores, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Pólo dos Açores, 9501-801 Ponta Delgada, Azores, Portugal
3
BIOPOLIS/CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Rua do Crasto, 765, 4485-684 Vairão, Vila do Conde, Portugal
4
Departamento de Biologia, Faculdade de Ciências e Tecnologia, Universidade dos Açores, 9501-801 Ponta Delgada, Açores, Portugal
5
UNESCO Chair—Land Within Sea: Biodiversity and Sustainability in Atlantic Islands, Universidade dos Açores, R. Mãe de Deus 13A, 9500-321 Ponta Delgada, Açores, Portugal
J. Mar. Sci. Eng. 2025, 13(11), 2024; https://doi.org/10.3390/jmse13112024
Submission received: 10 September 2025 / Revised: 10 October 2025 / Accepted: 17 October 2025 / Published: 22 October 2025
(This article belongs to the Special Issue Feature Review Papers in Geological Oceanography)
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Abstract

The fossil record of marine molluscs has been used as evidence to test marine island biogeography theories and to complement the evolutionary patterns of biodiversity and endemicity found in oceanic islands and archipelagos. Although long-distance dispersal patterns have been the subject of several studies, important questions may still be posed. For instance, are these processes random, or do they show distinct patterns? And are they restricted to “windows of opportunities”, or do they occur continuously? In the NE Atlantic, the dispersal of tropical species towards higher latitudes associated with the last interglacial period is a well-known phenomenon. However, the most probable dispersal route remains a matter of debate. To test these ideas, we used the Atlantic and Mediterranean last interglacial fossil records, and compared the present geographic distribution of shallow-water marine molluscs with that registered during the last interglacial episode, aiming to detect changes. Our results show that 27 species became extinct during the course of the last glacial episode, and that 55 marine mollusc species (43 gastropods and 12 bivalves) are ecostratigraphic indicators for the MIS 5e fossiliferous deposits in the Macaronesian archipelagos, the Atlantic coasts of Morocco, and the Mediterranean. Finally, we provide arguments for the timing of dispersal, which occurred during a restricted “window of opportunity” associated with the inception of the last interglacial, and for the most probable route of dispersal of the tropical species, most of them denoting a Cabo Verdean origin.

1. Introduction

Isolated oceanic islands and archipelagos are key places to study the biological evolution of shallow-water marine organisms [1,2,3,4]. Patterns of biogeography, biodiversity, and endemicity are commonly the result of such evolution [5,6,7,8,9,10,11,12]. One of the most important problems solved by the theory of marine island biogeography during recent decades was how marine species colonised faraway islands through long-distance dispersal. The issue of dispersal was pursued by many authors, but Rudolf Scheltema (1926–2019), working from the Woods Hole Oceanographic Institute in Massachusetts, was particularly successful in demonstrating the underlying patterns. For almost three decades, he explored the dispersal of marine organisms in detail [13,14,15]. As a result, he was able to correlate both the duration of the larval phase of marine gastropods with the dispersal ability of the species and with their geographic distributions, as well as the mode of larval development with the longevity of the species. In particular, Scheltema demonstrated that (1) marine mollusc gastropod species with planktotrophic larvae (i.e., larvae that feed during their more or less prolonged stay in the water column) have higher dispersal abilities and larger geographic ranges than non-planktotrophic species (including lecithotrophic larvae, i.e., larvae that do not feed during their short stay in the water column, and species with direct, intra-capsular development) [16,17,18,19,20,21,22]; and that (2) marine gastropod planktotrophic species generally possess longer lifespans (in a geological sense) compared to non-planktotrophic species [16,23,24,25].
Inspired by this framework, the author postulated [26] that relationships must be found between the mode of larval development, the dispersal process, the ecological zonation (i.e., the species’ usual bathymetric range), and the geographic range of the species. The resulting hypothesis suggests that, for congeneric non-planktotrophic gastropod species, those living in the intertidal zone should have wider geographic ranges and have speciated during a shorter time span than those living at greater depths [27]. These assumptions were later tested against both the fossil record [28,29] and molecular/phylogenetic data [30], and the results supported the theory.
Questions related to the long-distance dispersal of species are pertinent, and may still be posed. For example, are these processes random, or do they adhere to discernible patterns? Do they occur continuously, or are they more common during rare “windows of opportunities”? To test these ideas, the Atlantic and Mediterranean last interglacial fossil records are reviewed in order to detect range alterations on the geographic distribution of shallow-water marine molluscs, and to infer the most probable dispersal directions. Accordingly, this paper aims to address such issues, which are key to understanding the mechanisms responsible for the expansion and contraction of the geographic ranges of marine species during glacial/interglacial cycles, and to explain the patterns that emerge from migratory pathways across marine sweepstake routes that open during “windows of opportunity”, crossing former marine barriers in times of natural, global climate changes.

2. Materials and Methods

This study reviews and adds to previous work regarding the Marine Isotopic Stage 5e (MIS 5e, i.e., the warmest period of the last interglacial) deposits of the Macaronesia, the NW Atlantic coasts of Africa (from Senegal North to the Straits of Gibraltar), and the Mediterranean. Macaronesia is a geographic region that encompasses the Azores, Madeira, Selvagens, Canaries, and Cabo Verde archipelagos (Figure 1 and Figure 2). For the Macaronesian archipelagos, we reviewed the most relevant works by Meco [31,32], García-Talavera et al. [33,34], García-Talavera [35,36,37], Meco et al. [38,39,40,41], Callapez and Soares [42], García-Talavera and Sánchez-Pinto [43], Ávila et al. [44,45,46,47,48], Ávila [49], Cabero [50], Cabero et al. [51], Martín-González et al. [52,53], and Melo et al. [54,55]. For the African Atlantic coastal MIS 5e deposits, we critically reviewed the works of Lecointre [56,57,58], Ortlieb [59], Brebion [60], Brebion et al. [61], Weisrock et al. [62], Plaziat et al. [63], and Chakroun et al. [64]. Finally, the review benefitted from the compilation by [65] that summarises the mollusc record of the MIS 5e deposits of the Mediterranean, adding new MIS 5e mollusc records from recent important works [66,67].
Over the last ten years, more than 1500 primary sources (mainly research articles and monographs) and “grey literature” (about 30 Ph.D. and M.Sc. theses) have been consulted in the process of assembling databases on recent, and on MIS 5e shallow-water Atlantic (from the Arctic to the Antarctic) and Mediterranean mollusc species (gastropods and bivalves). For this work, two MIS 5e databases were formatted, both curated by the author, that include information on the geographic distribution and functional traits for a total of 1406 MIS 5e mollusc taxa: 887 gastropods and 519 bivalves. Information on the geographical distribution and functional traits of the present-day marine mollusc species was retrieved from two additional databases, also curated by the author: the first deals with the modern shallow-water (from the supralittoral down to 50 m depth) Atlantic and Mediterranean gastropods, with 5720 species [68], whereas the second database contains information on 1815 recent shallow-water (<100 m depth) Atlantic and Mediterranean bivalve species [69].
In order to detect possible species’ geographic range changes among the marine molluscs from the Macaronesian archipelagos, the Mediterranean, and along the Atlantic African coast, their present-day range was compared against their MIS 5e geographical distribution. This comparison facilitated the compilation of a checklist of MIS 5e molluscs that became extinct (Table 1), as well as of MIS 5e mollusc species/taxa that were extirpated, i.e., that locally disappeared from the Macaronesian archipelagos, NW Atlantic African coasts, and Mediterranean shores as a result of the last glacial episode (Supplementary Table S1). The cross-checking of mollusc species’ past and present-day geographic distributions also helped to determine the most probable sites of origin of the MIS 5e colonisers for each species, assuming that the most probable source of these colonisers was from the nearest archipelago or continental shores.
Regarding the NW Atlantic African coasts, it is necessary to exercise care, as doubts arise on correctly assigning some of the fossil mollusc species to the corresponding interglacials. Until recently, the major problem was the absence of fossil remains representing the large gastropod Thetystrombus latus (Gmelin, 1791) from the Atlantic coast of Morocco that would clarify this issue [59]. The solid sedimentary descriptions and stratigraphic analysis conducted by [70,71] provided such data for the Pleistocene (MIS 5e) coastal deposits of Rabat-Témara on the NW Atlantic coast of Morocco that, together with a precautionary use of older studies (mostly from the SW of Morocco, from Tan Tan beach to Tarfaya), allowed the compilation of a relevant set of MIS 5e Moroccan mollusc species to be included in our MIS 5e databases.
The data shown in Supplementary Materials Table S1 was additionally screened for the selection of MIS 5e ecostratigraphic indicators. As defined by [54], MIS 5e ecostratigraphic indicators are species that, cumulatively, fulfil the following conditions: (1) they are reported for the MIS 5e of the area in study (in our case, the Macaronesian archipelagos, the Atlantic coasts of Morocco and/or the Mediterranean); (2) these species do not occur today in the areas under study; and (3) these colonisers originate from a marine tropical region, i.e., an area located between 23.5° N and 23.5° S [72].
Finally, information on the functional traits of the ecostratigraphic indicator species was compiled, including data on the modes of larval development (planktotrophic/non-planktotrophic) and on the type of substrate (algae, gravel, rock, sand).

3. Results

From the 1406 MIS 5e species/taxa of molluscs (887 gastropods and 519 bivalves) reported from the entire Atlantic and Mediterranean, 27 species became extinct during the course of the last glacial episode (cf. Table 1): 18 gastropods and six bivalves in the Mediterranean; the gastropod Acanthina dontelei Garcia-Talavera & Sanchez-Pinto, 2002 in the Selvagens archipelago and in the Canaries; and one bivalve and one other gastropod species in the Canaries archipelago.
Besides the species that became extinct, another 137 MIS 5e species/taxa (38 Bivalvia, 99 Gastropoda) were extirpated from some of the areas where they occurred during the last interglacial (Supplementary Materials Table S1). Table 2 summarises the number of such species/taxa by area (for detailed information, please see Supplementary Materials Table S1). The Canary Islands, with 45 species/taxa, was the most affected archipelago, followed by the Mediterranean (35), Azores (27), Morocco (23), and Madeira (22). At the Selvagens archipelago, only two MIS 5e gastropods were extirpated and no longer live in the region: Isara nigra (Gmelin, 1791) and Patella ordinaria Mabille, 1888. The gastropod Gemophos viverratus (Kiener, 1834) was the only species that was extirpated from four different regions: the Azores and Madeira archipelagos, the Mediterranean, and the Atlantic coasts of Morocco. Three gastropod species—Conus ermineus Born, 1778, Conus ventricosus Bronn, 1831, and Linatella caudata (Gmelin, 1791)—were each extirpated from three regions: the first one from the Azores, Canaries, and Mediterranean; the second one from the Azores, Canaries, and Morocco; and the last one from the Azores, the Mediterranean, and Morocco (Supplementary Materials Table S1).
Table 3 displays the species/taxa of molluscs selected as MIS 5e ecostratigraphic indicators for the Azores, Madeira, Canaries, Cabo Verde, the Mediterranean, and Morocco, which are region-specific, based on information compiled in Supplementary Table S1, whereas Figure 3 and Figure 4 illustrate the dispersal directions of the MIS 5e mollusc ecostratigraphic indicators. Some species formerly included in the Gignoux “warm-water Senegalese fauna” [73], e.g., Monoplex trigonus (Gmelin, 1791) and Gemophos viverratus, were not considered ecostratigraphic indicators because their present geographic distribution is not restricted to tropical areas (both species occur today in the Canaries archipelago).
As expected, when all MIS 5e mollusc species/taxa that locally disappeared from the Macaronesian archipelagos, the Atlantic shores of Morocco, and/or from the Mediterranean coasts during the last glacial episode are included in the analysis, it becomes clear that most of the dispersal movements take place from tropical regions towards northern latitudes (Supplementary Materials Figures S1 and S2). Moreover, as shown in Figure 5, although the percentage of tropical gastropods that reached the Macaronesian archipelagos is similar, ranging from 45.8% in the Azores to 54.5% in the Canary Islands, it is much higher in the Mediterranean (77.8%) and much lower in the Atlantic Morocco region (30.4%). No tropical bivalves reached the Azores during the MIS 5e, only 20% reached Madeira, and about 60% reached Canaries; this contrasts with the MIS 5e bivalves that dispersed to the Atlantic coasts of Morocco and to the Mediterranean, all being tropical in provenience (Figure 5). As previously noted by [54], many longitudinal dispersal events were also detected, as well as range expansion towards lower latitudes, although fewer in number (Supplementary Materials Figures S1 and S2).

4. Discussion

4.1. Modes of Larval Development and Long-Distance Dispersal of Shallow-Water Marine Species: The Main Processes

Long-distance dispersal is one of the best-studied biogeographic processes by which marine species are able to break marine barriers to dispersal such as wide stretches of deep sea, strong upwelling regions, or large rivers and long estuaries [74,75,76,77]. To cross these barriers and succeed in reaching isolated islands/archipelagos, marine species use a variety of strategies including self-motility, foresy, rafting, and larval drift. Self-motility is well represented by the example of fishes [78,79] or marine mammals, such as the pinniped members of the Phocidae family [80]. Foresy is the term applied to hitchhiking on migratory shorebirds, attached to their feathers, bills, and legs [81,82,83] or within their digestive tract [84,85]. Floating rafts that enhance the dispersal of marine invertebrates is provided by algal masses [86,87,88,89,90,91,92,93,94], floating wood [95], or other natural objects such as floating rocks attached to kelps [96], ice [97], or floating pelagic crustacean barnacles [98], the carapaces of turtles [99], and even on pumice [100,101,102]. Perhaps the most common dispersal process occurs in the form of larvae [103,104], which are passively transported by trade and storm winds and by sea-surface currents.
The author’s database [68] includes information on the geographic distribution and 38 functional traits (longevity, shell composition, locomotion, life habit, diet, depth, maximum length, type of substrate) for 5720 recent Atlantic and Mediterranean shallow-water marine gastropod molluscs (shelled heterobranch gastropods are also included in the database). Out of these 5720 species, there are 161 gastropods (2.8%) that are amphi-Atlantic in geographic distribution. Of these 161, it was not possible to find information on the modes of larval development of 74 species; for the remaining species, the majority (52) are planktotrophic, 33 are non-planktotrophic, and two are brooding species [Capulus ungaricus (Linnaeus, 1758), and Littorina saxatilis (Olivi, 1792)]. This pattern is congruent with Scheltema rules [13,14,15]. Non-planktotrophic species are usually considered poor dispersers across sea barriers. Consequently, biologists have attributed the disjunct geographic distributions of non-planktotrophic species of marine gastropods to represent rare successful cases of long-distance dispersal on rafts of suitable, natural items by associated eggs, juveniles, or even small adult species [91]. Disjunct distributions are not restricted to molluscs, as examples are known from a variety of marine groups such as cnidarian hydrozoans [105], annelid polychaetes [106], echinoderms [107], and crustaceans [108,109].
Generally, for any given interval of time, it is expected that the number of planktotrophic gastropod species reaching isolated islands or archipelagos exceeds that of non-planktotrophic gastropod species. Moreover, as the geological lifespan of planktotrophic (p) gastropod species is usually longer (about 6–10 millions of years (Ma) on average; [110,111]) compared with that of non-planktotrophic (np) gastropod species (about 2–5 Ma on average; [110,111]), it is also expected that, for archipelagos/isolated islands older than 5 Ma, the number of planktotrophic species will be higher than that of non-planktotrophic gastropod species. However, our analysis, based on the database in [68], does not support these predictions, as for ten Atlantic archipelagos/islands studied (Table 4), only Bermuda clearly has a higher number of planktotrophic gastropods (150) in relation to the number of non-planktotrophic species (76). In the Azores, Madeira, Fernando de Noronha Island, and Trindade and Martim Vaz, the number of p-gastropods is similar to that of np-gastropods, and in the remaining archipelagos/islands, the proportion of np-gastropods is higher; therefore, other evolutionary factors must also be in action.
Several genera of marine gastropods portray relevant in situ adaptive radiations in the Atlantic islands (cf. Table 5), the most impressive being Euthria J. E. Gray, 1850 (92.9% of the 14 species are endemic); Mirpurina Ortea, Moro & Espinosa, 2019 (91.7% of the 12 species are endemic); Manzonia Brusina, 1870 (67.9% of the 28 species are endemic); Schwartziella G. Nevill 1881 (62.2% of the 37 species are endemic); Plesiocystiscus G. A. Coovert & H. K. Coovert, 1995 (46.7% of the 15 species are endemic); Chrysallida P. P. Carpenter, 1856 (45.5% of the 22 species are endemic); Fissurella Bruguière, 1789 (34.5% of the 29 species are endemic); Marginella Lamarck, 1799 (33.3% of the 30 species are endemic); Mitromorpha P. P. Carpenter, 1865 (33.3% of the 27 species are endemic); Crisilla Monterosato, 1917 (27.8% of the 36 species are endemic); Mitrella Risso, 1826 (25.0% of the 36 species are endemic); Conus Linnaeus, 1758 (21.1% of the 247 species reported from the Atlantic and Mediterranean are endemic to the Atlantic archipelagos and islands); Gibberula Swainson, 1840 (18.1% of the 116 species are endemic); and Odostomia J. Fleming, 1813 (14.3% of the 63 species are endemic). The Canaries is one of the oldest (25 Ma) and least isolated archipelagos in the Atlantic (presently just 98 km from the African continent). Together with Madeira and Selvagens, it forms the core of the Webbnesia biogeographic ecoregion [112], with many shared endemic and non-endemic species among these three archipelagos. Taken as a whole, these factors (age of the islands, insular littoral area, in situ speciation, and isolation) might explain the higher number of non-planktotrophic species in both Canaries and Selvagens. For further details, please see [10,11]. However, these arguments must be interpreted with some caution, as the number of gastropod species with unknown modes of larval development is still quite high, reaching 54.5% and 54.1% in São Tomé and Príncipe archipelago and Saint Helena Island, respectively (Table 4).

4.2. Relevance of the Fossil Record for Establishing the Times of Dispersal of Marine Species in the Macaronesian Archipelagos and in the Mediterranean

Both planktotrophic and non-planktotrophic marine species (these associated with rafts) are best able to cope with long-distance dispersal processes, thus reaching isolated islands and archipelagos. The fossil record helps to elucidate the time of arrival and past migratory pathways. In Macaronesia, fossiliferous sediments from the MIS 5e, with an attributed age of 129 to 116 ka [113], are well represented in all archipelagos: the Azores [10,28,35,42,44,45,47,48,114,115,116,117,118,119,120,121,122], Madeira [123], Selvagens [43], Canaries [31,32,33,36,38,39,40,50,51,52,53,124,125], and Cabo Verde [37,126].
The Mediterranean is the region with the highest number of MIS 5e fossil mollusc species/taxa reported (378) and is certainly one of the best-studied regions in the world for this time interval due to a very high publication effort relative to the marine biodiversity of the last interglacial period. This only highlights the significance of the MIS 5e fossil gastropods from the Azores, where over half of the present-day species were also recovered from the last interglacial deposits (55.6%) and, to a lesser degree, the Canaries and Atlantic Moroccan gastropod fossil record (26.0% and 26.8%, respectively; cf. Table 6). In contrast, the number of MIS 5e fossil bivalves of the Mediterranean (181 species/taxa; 48.4%) is impressive when compared with the number of shallow-water species of bivalves presently reported from the area (Table 6). These numbers also attest to the importance of continuous, several-decades-long studies, thus helping to lessen the Pleistocene taxonomic gap [48].
The marine Mollusca (especially the Gastropoda and the Bivalvia) is the best represented marine group in all Macaronesian archipelagos, followed by less common invertebrates (e.g., cnidarians, bryozoans, crustaceans, annelids, and echinoids), Chromista foraminifers, and vertebrate marine groups (e.g., bony fishes and cetaceans), as well as a few calcareous algal species. Quantitative data obtained from the fossil record of marine gastropods and bivalves derived from standardised 1 kg samples has been intensively used in the Macaronesia region for palaeoecological studies [46,47,127]. Additionally, the comparison between the insular faunas from the MIS 5e and those from the present day for each archipelago demonstrates the dispersal pathway patterns of many marine mollusc gastropods and bivalves from the continental shores of both the western and eastern Atlantic (i.e., from American, European, and African shores) to the Macaronesian archipelagos, as well as among the Macaronesian archipelagos. Moreover, these migrations took place during both interglacial [31,36,41,46,49,51,54,55,65,124,128] and glacial periods [127].
The MIS 5e is considered to have been the second-warmest interglacial period during the Quaternary, only behind the MIS 11c (424–397 ka; [129,130,131]. During the MIS 5e, the mean sea level was 6 to 9–10 m (MIS 5e) higher than today [132,133,134], with mean sea surface temperatures up to 3 °C higher than modern values [72,124,135,136]. Along the shores of the Macaronesian islands, the American and African coastlines, and in many coastal sites along the Mediterranean, marine terraces occur, ranging in altitude from 1 to over 10 m. Many of these erosional features include sediments rich in fossils from the last interglacial period (cf. Figure 2), and some of the molluscs found in these MIS 5e deposits in the Mediterranean and Atlantic Morocco and the Canaries, Madeira, and the Azores are warm-water species that do not occur today at these locations. This outcome suggests different climatic conditions during the last interglacial in comparison to those registered today.
Prior to the discovery by Milanković [137] of the orbital cycles that force the Quaternary climate, and based on the fossils collected and studied by the Italian geologist Domenico Lovisato (1842–1916), Arturo Issel described the “Tyrrhenian” stage of the Quaternary [138]. Originally called “Tirreno” [139], this stage refers to coastal marine fossiliferous sediments that were deposited during the last interglacial highstand at Cala Mosca (Gulf of Cagliari, Italy), now above present sea level. The fauna is typified by a warm-water malacofauna best represented by the iconic and large gastropod “Strombus bubonius” Lamarck, 1822, now accepted as Thetystrombus latus. Numerous authors reported on this warm-water fauna that characterises the MIS 5e deposits in various outcrops around the Mediterranean coasts, with the most relevant studies for this region being the Balearic Islands [140,141,142,143,144,145,146], Cyprus [147], Egypt [148], France [149], Greece [150,151], Israel [152], Italy [73,139,153,154,155,156,157,158,159,160,161,162,163,164,165,166,167,168,169,170], Lebanon [171], Southeast Spain [172,173,174], Tunisia [175,176,177,178,179,180,181,182,183], and Spain [184].
The MIS 5e interglacial period follows the glacial MIS 6, a period spanning from 191 to 130 ka [185]. The alternating interglacial and glacial periods, with warmer mean sea-surface temperature (SST) and humid climates for the former, and colder SST and arid climatic conditions for the latter, severely impacted the coastal marine biotas worldwide. One of the most important consequences was the effect that the changes in sea level had in the insular littoral area available for the settlement of shallow-water marine organisms. In insular habitats, and for over 80% of the islands [12], the littoral area reaches maximum values during interglacials and minimum values during glacials [10,11]. During the glacial Termination II (the small period of time between the end of the glacial MIS 6 and the inception of the MIS 5e), the sea level rose from about −130 m to +6 to +9 m in relation to the present sea level, and the end of this period is perceived by marine biogeographers as a “window of opportunity” for the range expansion of tropical species to higher latitudes [45,47,49]. In the NE Atlantic, some of these tropical species were able to enter into the Mediterranean via the Strait of Gibraltar, colonising its coastal habitats and establishing viable populations. The tropical species that migrated northwards have been informally called “Senegalese fauna”. Several authors have used this designation (e.g. [38,39,50,124,186,187,188] or a similar one (e.g., the well-known “couches a Strombes” used by Gignou [73,153]), with most of the authors referring to species that today thrive in a tropical area that ranges from Mauritania/Senegal south to Angola. In comparison, only a few authors [47,51] explicitly included species that today inhabit the Cabo Verde archipelago as members of the “Senegalese fauna”. Overall, these tropical Cabo Verdean/Senegalese species were subsequently extirpated from the northern Macaronesian archipelagos, the Mediterranean, and the Atlantic coasts of Morocco during the course of the last glacial episode.
Cabero [50] compiled a list of 23 “Senegalese” species (i.e., mollusc species associated with MIS 5e deposits in the Canary Islands and/or in the Mediterranean Sea, but that do not occur nowadays in either area), whereas Ávila and colleagues [47] identified 22 “southward ranging” species (sensu [39]), i.e., species associated with MIS 5e deposits in the Azores but that do not occur today in this archipelago. Later, Melo and colleagues [54] published a list of 23 ecostratigraphic indicators (10 bivalves, 13 gastropods) for the Macaronesian archipelagos. This list was criticised by Meco et al. [41], who rightly questioned the inclusion of some species. The opportunity has been taken to correct and update the list of MIS 5e ecostratigraphic indicators accepted as valid for the Macaronesian archipelagos, the Mediterranean, and Atlantic Morocco. The following species of bivalves were excluded from the list in [54] because of a present geographic distribution outside the tropical area (therefore not fulfilling the condition of being restricted to the tropics): Barbatia barbata (Linnaeus, 1758), Cardita rufescens Lamarck, 1819, Ostrea stentina Payeaudeau, 1826, Panopea sp., Pitar tumens (Gmelin, 1791), Senilia senilis (Linnaeus, 1758), and Spondylus gaederopus Linnaeus, 1758. In light of the current state of our knowledge, the author considers 12 to be the valid number of MIS 5e bivalve ecostratigraphic indicators for the Macaronesian archipelagos, the Atlantic shores of Morocco, and the Mediterranean coasts (see Table 7). Seven of these bivalve species (58.3%) are exclusively associated with hard grounds (rocks, shells, corals, coralline algae forming rhodoliths, gravel, pebbles or coarse sand), three bivalves (25.0%) are exclusively associated with soft, mobile bottoms (e.g., fine sand, sandy mud, or muddy sand), and two species (16.7%) are associated with mixed bottoms (cf. Table 7).
In contrast, of the 13 gastropod ecostratigraphic indicators for the MIS 5e of the Macaronesian archipelagos, only Cypraecassis testiculus (Linnaeus, 1758) and Turritella bicingulata Lamarck, 1822 are herein excluded from the list from [54], again because of a present geographic distribution outside the tropical area. Thus, the 43 gastropod ecostratigraphic indicators now accepted as valid for the MIS 5e include 22 species/taxa that are exclusive to the Macaronesian archipelagos, 3 species exclusive to the Atlantic coasts of Morocco, 13 species exclusive to the Mediterranean, and 5 species in common for both the Macaronesian archipelagos and the Mediterranean (Table 8). It was not possible to determine the type of substrate associated with 17 out of the 43 gastropod ecostratigraphic indicators. Of the remaining 26 species/taxa of gastropods, 13 (50.0%) are exclusively associated with hard bottoms, eight (30.8%) are exclusively associated with soft, mobile bottoms, and five species (19.2%) are associated with mixed bottoms (cf. Table 8).
The author’s databases [68,69] also allow for an interesting hypothesis to be put forward: the tropical gastropod species Alanbuella corrugata (Perry, 1811) (reported from the MIS 5e fossil record of the Canaries, Cabo Verde, Atlantic Morocco, and the Mediterranean), Monoplex trigonus (reported from the MIS 5e fossil record of the Canaries, Cabo Verde, and the Mediterranean), and Gemophos viverratus (reported from the MIS 5e fossil record of the Azores, Madeira, Canaries, Cabo Verde, Atlantic Morocco, and the Mediterranean) are tropical gastropod species in their present geographical distribution, except for the fact that they also occur today in the Canary Islands under a subtropical climate. It might have been the case that these three species, along with many others, expanded their geographical distribution during the last interglacial, but, in contrast with all other MIS 5e species, their Canary populations were able to cope with the global climate deterioration that followed the last interglacial and were not extirpated during the last glacial episode, with their existence in the archipelago becoming permanent.
As for the timing of the range expansion bringing these tropical mollusc species to higher latitudes, the author agrees with the mechanism proposed to break the previous marine barrier—the Canaries Current—between Cabo Verde and the northern Macaronesian archipelagos, and with the time for dispersal proposed by previous authors [11,45,47,49,54,55,116,189]. All of these authors agree that the detected northward range expansions of tropical fauna with a Cabo Verdean and west African origin occurred during a short “window of opportunity” just before the end of Termination II (the second Last Glacial termination, i.e., the period of time, c. 10 ka (140–130 ka), between the end of the MIS 6 glacial episode and the inception of the MIS 5e interglacial episode), or early in the last interglacial. Muhs et al. [189] clearly elucidated the process by suggesting that an orbitally forced mechanism triggered higher insolation in the early part of MIS 5e, and that this caused the northward shift of the intertropical convergence zone (ITCZ), complemented by weakened trade winds and reduced upwelling along the African coasts. According to [47], the average values for the summer insolation are inferred to have been about 11% higher than those of today. Ávila et al. [47] also stated that the oceanographic conditions prevalent during the earliest phase of the MIS 5e “moved the summer position of the Azores High to E of its present location, where it possibly persisted during the peak of the last interglacial, between the Azores and Iberia”, and that this shifted the biogeographic boundaries northward, a situation fully explored by [65] and its consequences explained in detail by these authors. To end this issue on the time of dispersal, it is necessary to explain the mechanism and the time for the southward migration of temperate and cold-water mollusc species during the MIS 5e that was captured by our data (see Supplementary Materials Figure S1C for a resume), first detected by [189] and later by [54]. We herein confirm their results and agree with the hypothesis put forward by the former authors that attributed this phenomenon to reduced insolation at the end of last interglacial, which caused a cascade of events that allowed for the southward dispersal of temperate and cold-water species: the southward displacement of the ITCZ, strengthening of the Canaries Current (as suggested by [65]), strengthening of the trade winds, and re-establishment of upwelling along the NW African coasts.

4.3. Main Routes of Dispersal in the East Atlantic During the MIS 5e

In the East Atlantic, two main routes are possible for the dispersal of a restricted number of tropical species that underwent latitudinal range expansions to higher latitudes in the Northern Hemisphere. This phenomenon occurred during the final phase of glacial Termination II/the early stages of the MIS 5e, as hypothesised by [11,41,45,47,49,65,189]. The first possible route of dispersal is an archipelagic route (either in a continuous stepping-stone fashion, or via a “jump-dispersal mode”, bypassing some of the archipelagos), with species originating in the Cabo Verde archipelago and migrating northward along the Macaronesian archipelagos, from Cabo Verde to the Canaries, Selvagens, and then to Madeira, with some species then entering the Mediterranean area, whereas other species were able to reach and colonise the Azores (see [31,35,37,39,42,43,45,46,47,50,52,53,54,55,123]). The second route of dispersal is along the NW continental shores of Africa, originating in the considered tropical areas (23.5° N and 23.5° S, i.e., from the south end of Western Sahara to Angola), and with adults creeping and/or larvae dispersing northward in times of a hypothesised reduction in the operation of the Mauritanian upwelling system and associated trade winds, finally entering and colonising the Mediterranean geographic area. Today, the Mauritania upwelling system, best expressed off Cap Blanc (Râs Nouâdhibou), constitutes a powerful marine biogeographic barrier for the dispersal of species and for the gene flow between populations of the same species inhabiting in the Canaries and Cabo Verde archipelagos, with an SST ranging from 14–15 °C in June to 21–22 °C in September–November [190]. The almost permanent regime of the Mauritania upwelling system results from the complex interactions between the coastal, shelf, and slope topography and the trade winds and currents that occur in the area [190]. The NNW-to-NNE trade winds drive the coastal upwelling off Mauritania, which takes place in the confluence zone where the northward Cabo Verde Current and Poleward Undercurrent meet the southward Canary Current [191,192]. These currents are deflected offshore and originate the SW directed Cabo Verde Frontal Zone, which displays a remarkable, giant Mauritanian chlorophyll filament [193]. As shown in Table 3 and in Supplementary Table S1, the longitudinal migration of species originating in the African shores towards similar Macaronesian archipelago latitudes also occur, an occurrence first detected by [54].
In order to choose between the two possible routes—the “Macaronesian archipelagic” or the “coastal African” route—detailed studies are necessary regarding the MIS 5e deposits of a large stretch of the African coast, from Senegal in the south, to the Atlantic Morocco Straits of Gibraltar in the north. The late Pleistocene fossiliferous deposits in the NW of Africa were studied by [56,57,58,59,60,61,62,63], just to name a few. A critical review of these works resulted in the compilation of a relevant set of 67 MIS 5e Moroccan mollusc species (23 bivalves and 44 gastropods) to be included in our MIS 5e databases. The comparison of the geographic distribution of the MIS 5e molluscs with their present-day occurrence yielded one species of bivalve and 22 species of gastropods that were extirpated from the MIS 5e Morocco Atlantic malacofauna during the last glacial episode. Although some gastropod species might have colonised the Atlantic coasts of Morocco from the Canary Islands and Madeira, or from Senegal and the Western Sahara, the majority seem to have most likely originated from the Mediterranean or from Iberian (Portugal) populations (cf. Supplementary Materials Figure S2). Thus, the geographic position of the Atlantic coasts of Morocco appears to render it a cornerstone position for the migration of marine species during the MIS 5e, with dispersal patterns much different to those registered at the Macaronesian archipelagos and the Mediterranean (cf. Supplementary Materials Figure S1C).
The functional traits of gastropods and bivalves may clarify the most important MIS 5e dispersal route for marine species. As shown in Table 7 and Table 8, most of the ecostratigraphic indicator bivalves and gastropods are associated with hard grounds (58.3% and 50%, respectively), with fewer bivalves and gastropods associated with soft, mobile bottoms (25% and 30.8%, respectively). In making this determination, one must take into account the following facts: (1) soft bottoms predominate in the long stretch of African coastline from Senegal to Atlantic Morocco; (2) hard grounds are extremely rare along the NW African coastline; (3) soft, mobile bottoms are available even in islands classified as “young” sensu Ávila et al. ([11], their Table 2), e.g., Terceira Island in the Azores, with just 0.4 Ma for its oldest subaerial age [194]; (4) hard substrata are common and sometimes even predominant in the islands that compose the Macaronesian archipelagos, in particular in those islands classified as “young” or “immature” sensu Ávila et al. [11]; (5) the existence of weakened trade winds and a reduced Mauritania upwelling system along the African coasts during the MIS 5e, as suggested by [189], implies that the SST should be lower in the continental coasts than those registered at similar latitudes in the southern Macaronesian archipelagos, in particular at the Canaries; (6) as suggested by [65], the transition zone between Tropical and Subtropical climatic zones was positioned at a higher latitude during the MIS 5e, resulting in the Canaries and the Cabo Verde archipelagos being located within the same tropical climatic zone; (7) the iconic ecostratigraphic indicator Thetystrombus latus is a thermophilic marine gastropod that is associated with sandy–muddy bottoms, favours waters of normal salinity and SSTs of around 25 °C [195], but with a temperature tolerance ranging from 15 °C to 31 °C [188]; (8) so far, no MIS 5e fossils of T. latus have been found in the NW African coasts between the Strait of Gibraltar and Senegal; and (9) MIS 5e fossils of T. latus are reported from Cabo Verde, the Canaries, and the Mediterranean.
When these previous data are coupled with the majority of the ecostratigraphic-indicator marine molluscs being associated with hard bottoms (cf. Table 7 and Table 8), the “Macaronesian archipelagic” route is most favourable in contrast to the less probable “coastal African” route. Although not with such strong and detailed arguments as those provided here, this “Macaronesian archipelagic” route was previously suggested by [45,47,49,54,55,65,128].

5. Conclusions

The last interglacial was one of the warmest interglacials during the last million years. Detailed knowledge of the MIS 5e fossil fauna from key sites such as the Macaronesian islands, the NW African coast, and the Mediterranean is crucial for a clear understanding of the various biogeographic processes and patterns that take place during glacial–interglacial transitions. This review suggests that the dispersal of tropical species to higher latitudes in the northern hemisphere, more specifically in the NE Atlantic, as registered in the last interglacial fossil record, is associated with a “window of opportunity”, i.e., a short period of time that comprises the ending of the glacial Termination II and/or the early stages of the MIS 5e. During this event, oceanographic and climatic conditions were different from those registered today, with a northern displaced ITCZ, a summer position of a weakened Azores High easterly displaced to a region between the Azores and Iberia, a weaker Canaries Current, reduced trade winds, and a reduced Mauritanian upwelling system. In addition to these conditions [196] also suggest that a persistent negative NAO (North Atlantic Oscillation) occurred at the end of glacial Termination II. Together, these conditions allowed tropical shallow-water marine molluscs to cross over the prevalent marine barrier formed by the Canaries Current and the Mauritanian upwelling system, thus allowing for the northward long-distance dispersal and geographic range expansion of Cabo Verdean marine species and, to a lesser degree, also of west African tropical species (mainly from Senegal, Mauritania, and Western Sahara). A thorough analysis was made of large datasets (5720 and 1815 recent gastropods and bivalves, respectively; and 887 and 519 MIS 5e gastropods and bivalves, respectively). The comparison of their geographic distributions during the MIS 5e and the present interglacial allowed for the detection of 27 species that became extinct due to the last glacial episode and the selection of 55 MIS 5e ecostratigraphic indicators specific for the Macaronesian archipelagos, the Atlantic coast of Morocco, and the Mediterranean. Finally, the fossil record data coupled with the information on the functional traits of the ecostratigraphic indicator species suggest a preferential archipelagic route along the Macaronesian archipelagos for the species that underwent long-distance dispersal processes, with some of them reaching latitudes as high as the Azores archipelago, whereas others, such as the large gastropod Thetystrombus latus, were able to reach and colonise the Mediterranean coasts.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/jmse13112024/s1: Figure S1: Total number (including ecostratigraphic and non-ecostratigraphic indicator species/taxa) and most probable origin of the MIS 5e mollusc species/taxa that underwent geographical range changes for the Azores, Madeira, Canaries, Cabo Verde, and the Mediterranean, according to Table S1. A: Gastropoda; B: Bivalvia. C: Sketch summarising the total number of molluscs (gastropods+bivalves) that underwent geographical range changes within the selected regions. Abbreviations as in Table S1; Figure S2: Total number (including ecostratigraphic and non-ecostratigraphic indicator species/taxa), and most probable origin of the MIS 5e mollusc species/taxa that underwent geographical range changes for Morocco Atlantic coasts, according to Table S1. A: Gastropoda; B: Bivalvia. Abbreviations as in Table S1; Table S1: MIS 5e mollusc species/taxa that locally disappeared from the Macaronesian archipelagos and/or from the Mediterranean coasts during the last glacial episode, with reference to their probable source/origin. The inferred most probable origin results from crosschecking information regarding the present and MIS 5e species’ geographic distribution. (*) corresponds to unlikely but still probable sites of origin of the colonisers. BIV: Bivalvia; GAS: Gastropoda; AZO: Azores; MAD: Madeira; SEL: Selvagens; CAN: Canaries; CAB: Cabo Verde; MED: Mediterranean; BER: Bermuda; MOR: Morocco; WES: Western Sahara; MAU: Mauritania; SEN: Senegal; IVO: Ivory Coast; POR: Portugal; CAR: Caribbean; VIR: Virginean biogeographic province (Atlantic shores of the USA, from Cape Cod (42° N) to Cape Hatteras, North Carolina (35°35′ N)).

Funding

SPA acknowledges his current FCT/2023.07418 CEEECIND research contract with BIOPOLIS (https://doi.org/10.54499/2023.07418.CEECIND/CP2845/CT0001). This work was supported by National Funds through FCT-Fundação para a Ciência e a Tecnologia in the scope of the project UID/50027-Rede de Investigação em Biodiversidade e Biologia Evolutiva. This work also benefitted from FEDER funds, through the Operational Program for Competitiveness Factors—COMPETE; National Funds through FCT (POCI-01–0145-FEDER-006821, UIDB/00153/2020, LA/P/0048/2020); and through the Regional Government of the Azores (M1.1.a/005/Funcionamento-C-/2016, CIBIO-A; M1.1.A/INFRAEST CIENT/A/001/2021; M3.3.B/ORG.R.C./005/2021, M3.3.B/ORG.R.C./008/2022/EDIÇÃO 1, M3.3.G/EXPEDIÇÕES CIENTÍFICAS/005/2022 and M3.3.G/EXPEDIÇÕES CIENTÍFICAS/004/2022). Finally, this work was also supported by FEDER funds (85%) and by funds from the Regional Government of the Azores (15%) through the Azores 2020 Operational Program under the VRPROTO project: Virtual Reality PROTOtype: the geological history of “Pedra- que-pica”: ACORES-01-0145-FEDER-000078, and by M1.1.A/INFRAEST CIENT/A/001/2021—Base de Dados da PaleoBiodiversidade da Macaronésia.

Data Availability Statement

The original contributions presented in this study are included in the article/Supplementary Materials. Further inquiries can be directed to the corresponding author.

Conflicts of Interest

The author declares no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Location of the Macaronesian archipelagos within the North Atlantic (insert) and geographic placement of the Macaronesian archipelagos.
Figure 1. Location of the Macaronesian archipelagos within the North Atlantic (insert) and geographic placement of the Macaronesian archipelagos.
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Figure 2. General views of MIS 5e outcrops in the Macaronesian archipelagos. (A): Vinha Velha (Santa Maria Island, Azores archipelago). (B): Zimbralinho (Porto Santo Island, Madeira archipelago). (C): Matas Blancas (Fuerteventura Island, Canaries archipelago), a famous outcrop renowned by the abundance of large specimens of the mollusc gastropod Thetystrombus latus (Gmelin, 1791). (D): Praia Lacacão (Boavista Island, Cabo Verde archipelago). All photos by the author.
Figure 2. General views of MIS 5e outcrops in the Macaronesian archipelagos. (A): Vinha Velha (Santa Maria Island, Azores archipelago). (B): Zimbralinho (Porto Santo Island, Madeira archipelago). (C): Matas Blancas (Fuerteventura Island, Canaries archipelago), a famous outcrop renowned by the abundance of large specimens of the mollusc gastropod Thetystrombus latus (Gmelin, 1791). (D): Praia Lacacão (Boavista Island, Cabo Verde archipelago). All photos by the author.
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Figure 3. The arrows indicate the number and most probable origin of the MIS 5e mollusc ecostratigraphic indicators for the Azores, Madeira, Canaries, Cabo Verde, and the Mediterranean, according to data in Table 3. (A): Gastropoda; (B): Bivalvia. Abbreviations as in Table 3.
Figure 3. The arrows indicate the number and most probable origin of the MIS 5e mollusc ecostratigraphic indicators for the Azores, Madeira, Canaries, Cabo Verde, and the Mediterranean, according to data in Table 3. (A): Gastropoda; (B): Bivalvia. Abbreviations as in Table 3.
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Figure 4. The arrows indicate the number and most probable origin of the MIS 5e mollusc ecostratigraphic indicators for Morocco, according to data in Table 3. (A): Gastropoda; (B): Bivalvia. Abbreviations as in Table 3.
Figure 4. The arrows indicate the number and most probable origin of the MIS 5e mollusc ecostratigraphic indicators for Morocco, according to data in Table 3. (A): Gastropoda; (B): Bivalvia. Abbreviations as in Table 3.
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Figure 5. Relative comparison of the warm water (tropical), temperate, cold water, and wide thermal range number of MIS 5e species (in percentage) for the Macaronesian archipelagos, the Atlantic coast of Morocco, and the Mediterranean. Abbreviations as in Table 3.
Figure 5. Relative comparison of the warm water (tropical), temperate, cold water, and wide thermal range number of MIS 5e species (in percentage) for the Macaronesian archipelagos, the Atlantic coast of Morocco, and the Mediterranean. Abbreviations as in Table 3.
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Table 1. List of MIS 5e species reported from the Macaronesian archipelagos and from the Mediterranean that became extinct (†) during the course of the last glacial episode. GAS: Gastropoda; BIV: Bivalvia; MED: Mediterranean; SEL: Selvagens archipelago; CAN: Canaries archipelago.
Table 1. List of MIS 5e species reported from the Macaronesian archipelagos and from the Mediterranean that became extinct (†) during the course of the last glacial episode. GAS: Gastropoda; BIV: Bivalvia; MED: Mediterranean; SEL: Selvagens archipelago; CAN: Canaries archipelago.
ClassMIS 5e Extinct SpeciesSiteGeological Timespan
EoceneOligoceneMiocenePliocenePleistoceneRecent
GASActeon bovettensis G. Seguenza, 1880 †MED 11
GASAlvania calliope Chirli & U. Linse, 2011 †MED 1
GASAlvania curta (Dujardin, 1837) †MED 1
GASAlvania mariae (A. d’Orbigny, 1852) †MED 1
GASAlvania unica Amati & Quaggiotto, 2019 †MED 1
GASCancilla alligata (Defrance, 1825) †MED 1
GASCapulus laevis (Bronn, 1831) †MED 1
GASCerithium diblasii †MED 1
GASClathromangelia clathrata (Serres, 1829) †MED 11
GASClathromangelia quadrillum (Dujardin, 1837) †MED 1
GASEpiscomitra fusiformis (Brocchi, 1814) †MED 1
GASDizoniopsis bilineata (M. Hörnes, 1848) †MED 11
GASFusinus rudis (R. A. Philippi, 1844) †MED 11
GASHelminthia triplicata (Brocchi, 1814) †MED 1
GASJujubinus bullula (P. Fischer, 1877) †MED 1
GASMetula deshayesi (Michelotti, 1847) †MED 1 1
GASOcinebrina scalaris (Brocchi, 1814) †MED 11
GASPetaloconchus intortus (Lamarck, 1818) †MED1 111
BIVEuropicardium multicostatum (Brocchi, 1814) †MED 1
BIVGlycymeris inflata (Brocchi, 1814) †MED 11
BIVMacoma obliqua (J. Sowerby, 1817) †MED 1
BIVMegaxinus transversus (Bronn, 1831) †MED 11
BIVPlicatula mytilina (R. A. Philippi, 1836) †MED 11
BIVPycnodonte squarrosa de Serres 1843 †MED 1 1
GASAcanthina dontelei Garcia-Talavera & Sanchez-Pinto, 2002 †SEL 1
GASAcanthina dontelei Garcia-Talavera & Sanchez-Pinto, 2002 †CAN 1
GASSpinucella plessisi (Lecointre, 1952) †CAN
BIVLucina columbella Lamarck, 1818 †CAN 1
Table 2. Total number of MIS 5e species/taxa that were extirpated, per region, during the course of the last glacial episode. For further details, please see Supplementary Materials Table S1.
Table 2. Total number of MIS 5e species/taxa that were extirpated, per region, during the course of the last glacial episode. For further details, please see Supplementary Materials Table S1.
AZOMADSELCANCABMORMED
Gastropoda221323262225
Bivalvia590136110
Total2722245122335
Table 3. The 55 species/taxa of molluscs selected as MIS 5e ecostratigraphic indicators for the Macaronesian archipelagos, Morocco Atlantic coasts, and the Mediterranean. The inferences made on the most probable origin of the colonising species results from crosschecking information regarding the present and MIS 5e species’ geographic distribution. (*) corresponds to unlikely but still probable sites of origin of the colonisers. BIV: Bivalvia; GAS: Gastropoda; AZO: Azores; MAD: Madeira; SEL: Selvagens; CAN: Canaries; CAB: Cabo Verde; MED: Mediterranean; BER: Bermuda; MOR: Morocco; WES: Western Sahara; MAU: Mauritania; SEN: Senegal; IVO: Ivory Coast; GAB: Gabon; POR: Portugal; CAR: Caribbean.
Table 3. The 55 species/taxa of molluscs selected as MIS 5e ecostratigraphic indicators for the Macaronesian archipelagos, Morocco Atlantic coasts, and the Mediterranean. The inferences made on the most probable origin of the colonising species results from crosschecking information regarding the present and MIS 5e species’ geographic distribution. (*) corresponds to unlikely but still probable sites of origin of the colonisers. BIV: Bivalvia; GAS: Gastropoda; AZO: Azores; MAD: Madeira; SEL: Selvagens; CAN: Canaries; CAB: Cabo Verde; MED: Mediterranean; BER: Bermuda; MOR: Morocco; WES: Western Sahara; MAU: Mauritania; SEN: Senegal; IVO: Ivory Coast; GAB: Gabon; POR: Portugal; CAR: Caribbean.
RegionClassSpecies/TaxaMost Probable Source/Origin
Range Expansion Towards Higher LatitudesRange Expansion Towards Lower LatitudesLongitudinal Range Expansion
AZOGASClaremontiella nodulosaCAB/SEN/BER
AZOGASConus ambiguusCAB/MAU
AZOGASConus ermineusCAB/WES/CAR *
AZOGASConus miruchaeCAB
AZOGASConus roeckeliCAB
AZOGASConus venulatusCAB
AZOGASZonaria pictaCAB/SEN
MADBIVCardium sp.MAU MOR *
MADGASAcliceratia carinataMAU
MADGASConus sp.CAB
MADGASThais nodosaCAB
MADGASZebina vitreaCAR
CANBIVAcar olivercoseliCAB/SEN
CANBIVBrachidontes puniceusCAB/MAU
CANBIVCardium sp.MAU MOR *
CANBIVChlamys sp.CAR
CANBIVCodakia sp.CARBER
CANBIVCtena eburneaCAB/SEN
CANGASCassis sp.CAB/BER *
CANGASCheilea equestrisCAB/BER *
CANGASClaremontiella nodulosaCAB/SEN/BER *
CANGASConus ermineusCAB WES
CANGASConus guinaicusSEN
CANGASCumia intertextaSEN
CANGASHarpa dorisCAB/SEN
CANGASHeliacus cylindricusCAB/BER *
CANGASHexaplex rosariumCAB/SEN
CANGASNerita senegalensisCAB/SEN
CANGASPrunum olivaeformeMAU
CANGASThais nodosaCAB
CANGASThetystrombus latusCAB/SEN
CANGASThylacodes arenariusCAB
CANGASVaughtia gruveliCAB WES
CANGASZonaria zonariaSEN
CABBIVSaccostrea cuccullataIVO SEN
CABGASFasciolaria sp. CAR
CABGASInermicosta inermicosta SEN
MEDBIVAcar olivercoseliCAB/SEN
MEDBIVAnadara geisseiCAB/MAU
MEDBIVArcopsis afraCAB/MOR
MEDBIVBrachidontes puniceusCAB/MAU
MEDBIVDendostrea cristataCAR
MEDBIVParvilucina crenellaCAR
MEDGASActeon maltzaniCAB/SEN
MEDGASActeon senegalensisMAU
MEDGASConus ermineusCAB/WES/CAR *
MEDGASConus guinaicusSEN
MEDGASEpitonium trochoidesMAU
MEDGASHexaplex rosariumCAB/SEN
MEDGASImbricariopsis carbonaceaCAB/SEN
MEDGASSimnia senegalensisCAB/SEN
MEDGASThetystrombus latusCAB/SEN
MEDGASTurbonilla kerstinaeCAB/MAU
MEDGASTurbonilla secernendaSEN
MEDGASXenophora senegalensisCAB/SEN
MEDGASZonaria petitianaMAU
MEDGASZonaria angolensisGAB
MORBIVCrassatina contrariaMAU
MORGASAnachis aurantiaSEN
MORGASCymbium gracileWES
MORGASHexaplex angularisSEN
Table 4. Total number of shallow-water (from the supralittoral down to 50 m depth) species of gastropods (# spp.), and percentage of species (% spp.) with unknown mode of larval development reported for each archipelago/island in the Atlantic Ocean. p: number of planktotrophic gastropod species; np: number of non-planktotrophic gastropod species. Data extracted from [68]. Age of the oldest island in each archipelago in millions of years (Ma), according to [10].
Table 4. Total number of shallow-water (from the supralittoral down to 50 m depth) species of gastropods (# spp.), and percentage of species (% spp.) with unknown mode of larval development reported for each archipelago/island in the Atlantic Ocean. p: number of planktotrophic gastropod species; np: number of non-planktotrophic gastropod species. Data extracted from [68]. Age of the oldest island in each archipelago in millions of years (Ma), according to [10].
Archipelago/Island# spp.pnp% spp. with
Unknown Mode of Larval Development		Age (Ma)
Azores224798128.66
Madeira3329810140.118.8
Selvagens156457324.429.5
Canaries69718021443.525
Cabo Verde52413014447.715.8
Bermuda4231507646.647
São Tomé and Príncipe369888054.59
Saint Helena Island3771054.115
Fernando de Noronha Island152626019.712.3
Trindade and Martim Vaz115504715.73.7
Table 5. Genera of Mediterranean and Atlantic gastropod molluscs with higher numbers of endemic species in the Atlantic archipelagos and islands. STP: São Tomé and Príncipe archipelago; STH: Saint Helena Island; ASC: Ascension Island; TRD: Trindade and Martim Vaz archipelago; ASP: São Pedro and São Paulo archipelago; FNO: Fernando de Noronha archipelago; FAL: Falkland Islands. For other abbreviations, see the legend of Table 3.
Table 5. Genera of Mediterranean and Atlantic gastropod molluscs with higher numbers of endemic species in the Atlantic archipelagos and islands. STP: São Tomé and Príncipe archipelago; STH: Saint Helena Island; ASC: Ascension Island; TRD: Trindade and Martim Vaz archipelago; ASP: São Pedro and São Paulo archipelago; FNO: Fernando de Noronha archipelago; FAL: Falkland Islands. For other abbreviations, see the legend of Table 3.
GenusTotal Number of SpeciesArchipelagos/Islands% of Endemic Insular Species
AZOMADSELCANCABBERSTPSTHASCTRDASPFNOFAL
Alvania Risso, 18261367 15823 19.1
Ammonicera Vayssière, 189316 2 12.5
Anachis H. Adams & A. Adams, 185321 2 1 14.3
Barleeia W. Clark, 185312 3 1 33.3
Bittium Leach, 1847151 5 40.0
Botryphallus Ponder, 199031 1 66.7
Caecum J. Fleming, 1813832 236 1 16.9
Chrysallida P. P. Carpenter, 185622 10 45.5
Conus Linnaeus, 1758247 149 1 1 21.1
Coralliophila H. Adams & A. Adams, 185323 1 11 13.0
Crisilla Monterosato, 191736 46 27.8
Diodora J. E. Gray, 182140 12 1 10.0
Eatonina Thiele, 191211 2 18.2
Euthria J. E. Gray, 185014 13 92.9
Fissurella Bruguière, 178929 6 11234.5
Gibberula Swainson, 18401161 103 4 3 18.1
Gibbula Risso, 1826201 4 25.0
Hastula H. Adams & A. Adams, 185322 12 1 18.2
Jujubinus Monterosato, 1884231 31 21.7
Manzonia Brusina, 18702821177 1 67.9
Marginella Lamarck, 179930 10 33.3
Metaxia Monterosato, 188415 2 1 20.0
Mirpurina Ortea, Moro & Espinosa, 201912 11 91.7
Mitrella Risso, 182636 4 5 25.0
Mitromorpha P. P. Carpenter, 1865271 4 4 33.3
Muricopsis Bucquoy & Dautzenberg, 188219 4 21.1
Odostomia J. Fleming, 18136321 3 12 14.3
Onoba H. Adams & A. Adams, 18523511 1 111.4
Parviturbo Pilsbry & T. L. McGinty, 1945141 2 21.4
Patella Linnaeus, 175813 1 11 23.1
Phorcus Risso, 182610 1 1 20.0
Plesiocystiscus G. A. Coovert & H. K. Coovert, 199515 1 33 46.7
Pradoxa F. Fernandes & Rolán, 19934 4 100.0
Pseudoscilla O. Boettger, 19025 1 1 40.0
Putzeysia Sulliotti, 18893 1 2 100.0
Pyrgocythara Woodring, 192810 1 10.0
Rissoa Desmarest, 1814 371 31 13.5
Rissoella J.E. Gray, 1847313 11 16.1
Schwartziella G. Nevill 1881 37 19 31 62.2
Volvarina Hinds, 184426611 12611 18.6
TOTAL 2553571748481246113
Table 6. Total number of species of Gastropoda and Bivalvia reported from the MIS 5e deposits of the Macaronesian archipelagos, the Atlantic coasts of Morocco, and the Mediterranean, and comparison with the present fauna in the same sites (in %). Abbreviations as in Table 3.
Table 6. Total number of species of Gastropoda and Bivalvia reported from the MIS 5e deposits of the Macaronesian archipelagos, the Atlantic coasts of Morocco, and the Mediterranean, and comparison with the present fauna in the same sites (in %). Abbreviations as in Table 3.
GastropodaAZOMADSELCANCABMORMED
MIS 5e12553181825344378
Present-day2253331566995281641069
MIS 5e/Present-day (%)55.615.911.526.010.026.835.4
BivalviaAZOMADSELCANCABMORMED
MIS 5e28292793022181
Present-day15013539233130287374
MIS 5e/Present-day (%)18.721.55.133.923.17.748.4
Table 7. List of the selected MIS 5e bivalve ecostratigraphic indicators for the Macaronesian archipelagos, Atlantic Morocco, and the Mediterranean, and respective type of substrate. Abbreviations as in Table 3.
Table 7. List of the selected MIS 5e bivalve ecostratigraphic indicators for the Macaronesian archipelagos, Atlantic Morocco, and the Mediterranean, and respective type of substrate. Abbreviations as in Table 3.
Bivalve Species/TaxaMIS 5e Geographic Range Expansion to:SUBSTRATE
Hard Grounds Gravel, PebblesCoarse SandFine SandSandy Mud-Muddy Sand
Acar olivercoseliCAN/MED1
Anadara geisseiMED 1
Arcopsis afraMED1
Brachidontes puniceusCAN/MED1
Cardium sp.MAD/CAN 11
Chlamys sp.CAN11 1
Codakia sp.CAN 11
Crassatina contrariaMOR
Ctena eburneaCAN 11
Dendostrea cristataMED1
Parvilucina crenellaMED 11
Saccostrea cuccullataCAB11
Table 8. List of the selected MIS 5e gastropod ecostratigraphic indicators for the Macaronesian archipelagos, Atlantic Morocco, and the Mediterranean, and respective type of substrate. Abbreviations as in Table 3.
Table 8. List of the selected MIS 5e gastropod ecostratigraphic indicators for the Macaronesian archipelagos, Atlantic Morocco, and the Mediterranean, and respective type of substrate. Abbreviations as in Table 3.
Gastropod Species/TaxaMIS 5e Geographic Range Expansion to:SUBSTRATE
Hard GroundGravel, PebblesCoarse SandFine SandSandy Mud-Muddy Sand
Acliceratia carinataMAD
Acteon maltzaniMED
Acteon senegalensisMED
Anachis aurantiaMOR1
Aporrhais senegalensisMED
Cassis sp.CAN1 11
Cheilea equestresCAN1
Claremontiella nodulosaAZO/CAN1
Conus sp.MAD1
Conus ambiguusAZO11
Conus ermineusAZO/CAN/MED1 1
Conus guinaicusCAN/MED
Conus miruchaeAZO
Conus roeckeliAZO
Conus venulatusAZO
Cumia intertextaCAN
Cymbium gracileMOR 1
Epitonium trochoidesMED 1
Fasciolaria sp.CAB
Gemophos viverratusAZO/MAD/MED1
Harpa dorisCAN 1
Heliacus cylindricusCAN 111
Hexaplex angularisMOR1
Hexaplex rosariumCAN/MED1
Imbricariopsis carbonaceaMED
Inermicosta inermicostaCAB1
Monoplex trigonusMED1 1
Nerita senegalensisCAN1
Prunum olivaeformeCAN
Simnia senegalensisMED1
Terebra corrugataMED 1
Thais nodosaMAD/CAN1
Thetystrombus latusCAN/MED1 1
Thylacodes arenariusCAN
Turbonilla kerstinaeMED
Turbonilla secernendaMED
Vaughtia gruveliCAN
Xenophora senegalensisMED 1
Zebina vítreaMAD
Zonaria angolensisMED 1
Zonaria petitianaMED 1
Zonaria pictaAZO 1
Zonaria zonariaCAN1
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Ávila, S.P. Distributional Range Shifts Caused by Glacial–Interglacial Cycles: A Review on Timing, Main Processes, and Patterns of Late Pleistocene Marine Dispersal by Invertebrates in the NE Atlantic. J. Mar. Sci. Eng. 2025, 13, 2024. https://doi.org/10.3390/jmse13112024

AMA Style

Ávila SP. Distributional Range Shifts Caused by Glacial–Interglacial Cycles: A Review on Timing, Main Processes, and Patterns of Late Pleistocene Marine Dispersal by Invertebrates in the NE Atlantic. Journal of Marine Science and Engineering. 2025; 13(11):2024. https://doi.org/10.3390/jmse13112024

Chicago/Turabian Style

Ávila, Sérgio P. 2025. "Distributional Range Shifts Caused by Glacial–Interglacial Cycles: A Review on Timing, Main Processes, and Patterns of Late Pleistocene Marine Dispersal by Invertebrates in the NE Atlantic" Journal of Marine Science and Engineering 13, no. 11: 2024. https://doi.org/10.3390/jmse13112024

APA Style

Ávila, S. P. (2025). Distributional Range Shifts Caused by Glacial–Interglacial Cycles: A Review on Timing, Main Processes, and Patterns of Late Pleistocene Marine Dispersal by Invertebrates in the NE Atlantic. Journal of Marine Science and Engineering, 13(11), 2024. https://doi.org/10.3390/jmse13112024

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