A revision of the Encarsia mexicana species-group (=Dirphys Howard) (Hymenoptera: Aphelinidae), gregarious endoparasitoids of whiteflies (Hemiptera: Aleyrodidae) in the Neotropical Region

Simple Summary Encarsia (Family Aphelinidae) is a genus of minute parasitoid wasps that target a diversity of agricultural pest insects including whiteflies and armoured scale insects. Since the genus was described in 1878, 450+ species of Encarsia have been described. Historically, it has been difficult to provide a subgeneric classification for the species of Encarsia. As a result, researchers have divided the group into numerous informal species-groups. Our work uses an alignment of ribosomal DNA sequences (sequences that code for ribosomal RNA) to construct a phylogenetic tree that shows that species of the genus Dirphys are correctly placed within Encarsia. With these results, we establish the Encarsia mexicana species-group (for the six species previously placed in Dirphys) and describe 14 new species. We also briefly discuss morphological characters that may correspond to the relationships recovered in the molecular phylogeny. With this information, we can better understand the patterns of evolution which brought about the present diversity within Encarsia and provide a more accurate classification of the genus. Abstract The genus Dirphys Howard 1914 syn. n. is synonymized with Encarsia Förster, and treated as a species-group of Encarsia, referred to henceforth as the Encarsia mexicana species-group. The monophyly of Encarsia is discussed in relation to Dirphys. The new synonymy is based on phylogenetic analyses of the nuclear ribosomal 28S-D2 gene region (43 taxa, 510 bp). The Encarsia mexicana species-group is recovered as strongly monophyletic within Encarsia. All species of the Encarsia mexicana species-group are revised. The group includes six previously described species, and fourteen newly described species. All species are described (or redescribed) and illustrated. Detailed distributional data, and, where available, plant associate and host records are provided for all species. Encarsia myartsevae Kresslein and Polaszek nom. nov. is here proposed as a replacement name for Encarsia mexicana Myartseva, now preoccupied by Encarsia mexicana (Howard). A dichotomous identification key, supplemented by an online multiple-entry key, is provided for all species.


Introduction
The genus Dirphys was initially described by Howard [1] for Mesidia mexicana Howard [2]. Where known, species in this genus are primary endoparasitoids of Aleyrodidae [3][4][5] and are gregarious, with up to 16 developing in a single host [3,4]. This behavior is unknown 2. 3

. DNA Extraction, Amplification, and Sequencing
Genomic DNA was extracted from single, whole specimens using a non-destructive genomic DNA extraction protocol developed by Chao-Dong Zhu, John Noyes, and others at the Natural History Museum, London [17]. Occasionally 2-3 specimens were extracted together when known to be conspecific.
Specimens were softened in 70% ethanol (to reduce potential damage during subsequent steps) at room temperature for a minimum of 2 h. Seventy percent ethanol was removed carefully by pipette and specimens were allowed to air-dry briefly. DNA was extracted using the Qiagen DNeasy Blood and Tissue Kit (250) 69506 (Qiagen, Hilden, Germany). Specimens were immersed in 180 µL of Lysis Buffer ATL, premixed with 20 µL Proteinase K and incubated at 55 • C overnight (8 h minimum) with no mixing, taking care that the specimen was submerged/floating in the buffer and not adhered to the side of the tube.
After digestion, the lysis buffer was carefully removed by pipette into a clean 1.5 mL microfuge tube. The specimen was immediately washed by adding 500 µL distilled water for a minimum of 30 min, then replaced with 500 µL 70% ethanol for a minimum of 30 min, then finally stored in 100% ethanol until slide-mounted in Canada balsam.
DNA was extracted from the lysis buffer using the Qiagen QUIA quick PCR Purification Kit (250) 28106 following the protocol: 'Isolation of total DNA from Animal Tissues' (step 3 onwards). Standard PCR reactions were then carried out in a thermal cycler using 2.0 µL DNA extract, Taq buffer (1.5 mM MgCl2), 1.5 U Taq polymerase (Roche), 10 nmol dNTPs (Amersham Pharmacia Biotech; APB) and 20 mol of each primer at the Natural History Museum's DNA sequencing facility. The D2 region of 28S rDNA was amplified using the following primers: 28SFW 5 -AGTACCGTGAGGGAAAGTTG-3 28SRev 5 -TTGGTCCGTGTTTCAAGACGG 3 PCR conditions were as follows: an initial denaturation of 94 • C for 3 min, then 35 cycles of denaturation at 94 • C for 1 min, annealing at 50 • C for 1 min, and extension at 72 • C for 2 min, followed by a final extension at 75 • C for 10 min, then samples were held at 4 • C until they could be analyzed. PCR products were run on a 1% agarose gel to confirm PCR success (clean bands of the expected size), then the remaining products were cleaned and sequenced. Removal of dye terminators was done by ethanol precipitation prior to sequencing. Specimens were imaged using a Leitz Dialux 20EB (Wetzlar, Germany) compound microscope using Nomarski Differential Interference Contrast illumination (DIC) and photographed with a MicroPublisher 5.0 RTV (Qiagen, Hilden, Germany) camera. Additional images (claval sensorial area; mandibles) were imaged with an Olympus BX63 microscope (Olympus, Tokyo, Japan) also utilizing DIC. Scanned sections were stacked and combined using Synoptics AutoMontage Pro ® ver. 5.03 software (Leitz Dialux images) and Helicon Focus software (Olympus BX63 images). The final images were edited with Adobe Photoshop CC ® .

DNA Extraction, Amplification, and Sequencing
Genomic DNA was extracted from single, whole specimens using a non-destructive genomic DNA extraction protocol developed by Chao-Dong Zhu, John Noyes, and others at the Natural History Museum, London [17]. Occasionally 2-3 specimens were extracted together when known to be conspecific.
Specimens were softened in 70% ethanol (to reduce potential damage during subsequent steps) at room temperature for a minimum of 2 h. Seventy percent ethanol was removed carefully by pipette and specimens were allowed to air-dry briefly. DNA was extracted using the Qiagen DNeasy Blood and Tissue Kit (250) #69506 (Qiagen, Hilden, Germany). Specimens were immersed in 180 µL of Lysis Buffer ATL, premixed with 20 µL Proteinase K and incubated at 55 °C overnight (8 h minimum) with no mixing, taking care that the specimen was submerged/floating in the buffer and not adhered to the side of the After digestion, the lysis buffer was carefully removed by pipette into a clean 1.5 mL microfuge tube. The specimen was immediately washed by adding 500 µL distilled water for a minimum of 30 min, then replaced with 500 µL 70% ethanol for a minimum of 30 min, then finally stored in 100% ethanol until slide-mounted in Canada balsam. The DNA analyzer system ABI PRISM 3730 and 377 DNA sequencer were used, the samples were loaded onto the system's vertical polyacrylamide gel where they underwent electrophoresis, laser detection and computer analysis. Sequence editing and alignment were performed using Sequencher TM 4.8 (Gene Codes Corporation, Ann Arbor, USA) on a Macintosh computer. The resulting molecular dataset includes 18 sequences representing 13 species. Sequences have been deposited in the GenBank database under accession numbers OQ683545-OQ683576 (Tables 1 and 2).

Phylogenetic Analyses
Captured sequences were combined with previously published sequence data (Table 2) and aligned using the E-INS-I algorithm in MAFFT v7.490 [18]. Ten independent iterations of maximum likelihood were reconstructed using IQ-TREE version 2.0.7 [19], implementing a General Time Reversible model with invariant sites and gamma distributed rate variation (-m GTR + I + G). Bootstrap support was estimated from 1000 bootstrap trees constructed using ultrafast bootstrapping (-b 1000) [20]. Outgroups are comprised of a broad range of Encarsia species, a diversity of recognized species-groups, as well as Coccophagus Westwood (Aphelinidae: Coccophaginae) and Aphytis Howard (Aphelinidae: Aphelininae).

Nomenclatural Acts
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved, and the associated information can be viewed through any standard web browser by appending the LSID to the prefix "http://zoobank.org/ (accessed on 6 June 2023)". The LSID for this publication is urn:lsid:zoobank.org:pub:2CE58923-A39A-412A-896E-DCFD4CC01FD7. The electronic edition of this work was published in a journal with an ISSN and has been archived and is available from the following digital repositories: PubMed Central, LOCKSS.

Phylogenetic Analysis of Molecular Data
A maximum likelihood tree was constructed from partial sequences of 28S D2 ribosomal DNA of 13 species (from 18 specimens) and 23 outgroup taxa ( Figure 2). The Encarsia mexicana species-group was recovered as a strongly supported clade within Encarsia. Encarsia dichaeta forms the sister clade to all remaining E. mexicana-group species. The Encarsia mexicana species-group was not placed sister to the noyesi species-group; however, backbone support in the recovered phylogeny is insufficient to confidently resolve interand intra-species-group relationships.

Taxonomy of the Encarsia mexicana Species-Group
Encarsia mexicana species-group = Dirphys Howard, 1914 Etymology of Dirphys. Dirphys (∆ιρϕυς) is a Greek feminine noun. Hence the modification by Hayat of Howard's (1914) combination Dirphys mexicana (Howard) to Dirphys mexicanus (Howard) was an unjustified emendation [21]. Hayat attributed the new combination to Howard [1], but this is not the case. Diagnosis: head dorsally transverse. Frontovertex at narrowest wider than dorsal eye width. Facial lines evident, often broadly expanded; mediofrontal and transfacial lines developed. Eyes with evident setae. Mandibles usually with two teeth and a truncation ( Figure 10A), the truncation sometimes reduced, and the teeth often strongly developed so mandibles appear to have only two teeth ( Figure 22A). A bidentate upper tooth may be present in addition to the well-developed ventral tooth ( Figure 4A).
Maxillary palps two-segmented. Antenna eight-segmented in both sexes, antennal formula variable (1, 1, 3, 3 or 1, 1, 0, 6), claval sensorial complex present ( Figure 17B) or absent, suture between F5 perpendicular or oblique. Pronotum medially membranous. Mesoscutum with more than 20 setae. Side lobes divided ( Figure 23E). Axillae large, strongly projecting forwards and separated medially by less than the maximum length of one axilla. Each axilla usually with a single seta, in E. dichaeta with two setae ( Figure 11E). Thoracic sculpture aciculate, longitudinal, transverse or a combination of these types, never reticulate, always strongly rugose. Mesoscutellar sensilla close together, separated by about the width of one sensillum. Fore wings with two large setae on the submarginal vein, plus a variable number of smaller setae at the distal end of the submarginal vein. Linea calva present or absent. Mid basitarsus with a variable number of robust, spine-like setae, tarsi five-segmented.
Remarks: The Encarsia mexicana species-group ( Figure 3) is restricted to the Neotropical zone, with species reaching as far south as the State of Bahia (Brazil), and as far north as southern Mexico. Encarsia dichaeta forms the sister clade to all remaining E. mexicana-group species. The Encarsia mexicana species-group was not placed sister to the noyesi species-group; however, backbone support in the recovered phylogeny is insufficient to confidently resolve interand intra-species-group relationships.

Taxonomy of the Encarsia Mexicana Species-Group
Encarsia mexicana species-group = Dirphys Howard, 1914 Etymology of Dirphys. Dirphys (Διρφυς) is a Greek feminine noun. Hence the modification by Hayat of Howard's (1914) combination Dirphys mexicana (Howard) to Dirphys mexicanus (Howard) was an unjustified emendation [21]. Hayat attributed the new combination to Howard [1], but this is not the case.
Diagnosis: head dorsally transverse. Frontovertex at narrowest wider than dorsal eye width. Facial lines evident, often broadly expanded; mediofrontal and transfacial lines developed. Eyes with evident setae. Mandibles usually with two teeth and a truncation ( Figure 10A), the truncation sometimes reduced, and the teeth often strongly developed so mandibles appear to have only two teeth ( Figure 22A). A bidentate upper tooth may be present in addition to the well-developed ventral tooth ( Figure 4A). Maxillary palps two-segmented. Antenna eight-segmented in both sexes, antennal formula variable (1, 1, 3, 3 or 1, 1, 0, 6), claval sensorial complex present ( Figure 17B) or absent, suture between F5 perpendicular or oblique. Pronotum medially membranous. Mesoscutum with more than 20 setae. Side lobes divided ( Figure 23E). Axillae large, strongly projecting forwards and separated medially by less than the maximum length of one axilla. Each axilla usually with a single seta, in E. dichaeta with two setae ( Figure 11E).  by about the width of one sensillum. Fore wings with two large setae on the submarginal vein, plus a variable number of smaller setae at the distal end of the submarginal vein. Linea calva present or absent. Mid basitarsus with a variable number of robust, spine-like setae, tarsi five-segmented.
Remarks: The Encarsia mexicana species-group ( Figure 3) is restricted to the Neotropical zone, with species reaching as far south as the State of Bahia (Brazil), and as far north as southern Mexico.      Remarks: T7 is extremely extended, covering the ovipositor. Encarsia acusa appears to be most closely related to E. inbioa and E. svetlana, but is easily distinguished from those (and all other) species by the extremely long ovipositor and T7. DNA sequences from holotype deposited under GenBank accession numbers: OQ683554. Etymology. From "acus" Latin for needle or pin, referring to the elongated T7. Remarks: T7 extremely extended, covering the ovipositor. Encarsia acusa appears to be most closely related to E. inbioa and E. svetlana, but is easily distinguished from those (and all other) species by the extremely long ovipositor and T7. DNA sequences from holotype deposited under GenBank accession number: OQ683554. Etymology. From "acus" Latin for needle or pin, referring to the elongated T7.

Encarsia aisha
Polaszek and Hernández-Suárez sp. n. Remarks: Encarsia aisha is morphologically very similar to E. marynoyesae in many respects (though distant to it based on DNA). The species can be distinguished from E. marynoyesae by the second valvifers almost 2x (1.8) the third valvulae; while they are 1.5x as long in E. aisha. In E. marynoyesaei the clava is well over 2x the length of the funicle; in E. aisha it is less than 2x as long. DNA sequence from holotype and paratype (pooled extraction) deposited under GenBank accession number: OQ683562.
Etymology: Named for Aisha, daughter of the second author (EHS), and sister to Noora; see E. noora, below.
Distribution Remarks: Encarsia aphania presents a unique combination of characters and appears to have no very close relatives. Morphologically it is closest to E. larensis but is easily distinguishable by the much longer third valvulae relative to the second valvifers (compare Figures 5F and 15F). DNA sequences were obtained from two specimens from Belize (type locality) and Costa Rica, Puntarenas; deposited under GenBank accession numbers: OQ683546, OQ683545.

Encarsia avida
Polaszek and Hernández-Suárez sp. n. Remarks: Encarsia avida appears morphologically close to E. acusa with which it shares the color pattern (mesoscutellum anteriorly dark) and wing and antennal morphology.
The ovipositor in E. acusa is longer (1.8x mid tibia; 1.6x in E. avida). The most easily appreciated difference is in the sculpture of the frons: E. avida has scattered, shallow horizontal grooves ( Figure 7A) while E. acusa has very dense horizontal grooves ( Figure 4A  Remarks: Encarsia avida appears morphologically close to E. acusa with which it shares the color pattern (mesoscutellum anteriorly dark) and wing and antennal morphology.
The ovipositor in E. acusa is longer (1.8x mid tibia; 1.6x in E. avida). The most easily appreciated difference is in the sculpture of the frons: E. avida has scattered, shallow horizontal grooves ( Figure 7A) while E. acusa has very dense horizontal grooves ( Figure 4A). A similar difference in sculpture is evident on the lateral face. The two species are wellseparated based on DNA (Figure 1) with E. avida coming out as sister to E. aphania with high support (95%). DNA sequence from holotype deposited under GenBank accession number: OQ683547.
( Figure 8A-F) Female. Color: Antennae brown. Head dark brown. Mesosoma uniformly dark brown. Legs yellow with mid and hind femora, coxae, and anterior half of tibiae brown, fore leg femora, coxae, and tibiae dark, all tarsi pale. Wings infuscate below marginal vein, submarginal and marginal veins dark, stigmal vein paler.  Remarks: Encarsia catula shares aspects of its morphology with E. marynoyesae, but can be distinguished by having fewer than 30 setae on the mesoscutum, and third valvulae more than 1 2 the length of second valvifers (less than 1 2 as long in E. marynoyesae). The two species are relatively close based on DNA (Figure 1). DNA sequence from the holotype is deposited under GenBank accession number: OQ683547.
( Figure 10A-F) Dirphys diablejo Polaszek and Hayat, 1992: 189 Female. Unknown. This species is known only from the holotype.   Figure 10A) with two large pointed teeth and a truncation; maxillary palps two-segmented. Mid-lobe of mesoscutum ( Figure 10E) with more than 60 setae; each lateral lobe with one seta; each axilla with one seta; scutellum with four setae and two vestigial setal bases. Sculpture of mesoscutum transverse. Fore wing ( Figure 10C Remarks: For the purposes of the identification key, we have assumed that the (unknown) female of E. diablejo shares the wing and mesosomal sculpture characters with the male; the combination of which is unique in the Encarsia mexicana species-group. 3 Figure 11D) equal to corresponding basitarsus. Metasomal terga T1-T7 with 0, 1 + 1, 1 + 1, 1 + 1, 1 + 2 + 1, 1 + 2 + 1 and 12 setae, respectively. T7 ( Figure 11F Remarks: There are some colour differences between the Costa Rican specimens and those from Brazil, the latter having the metasoma distally paler. Further studies on fresh material, in particular DNA sequencing, will be needed to confirm their status. DNA sequences were obtained from the holotype and five paratypes, deposited under GenBank accession numbers: OQ683550, OQ683552, OQ683551 and OQ683549 (three paratype specimens were pooled for extraction).
Etymology. "dichaeta" refers to the two setae on each axilla, unique for the genus.  Remarks: Encarsia encantadora is morphologically closest to E. erwini, differing from that species mainly in having the third valvulae longer than the second valvifers. The fore wing is also broader in E. encantadora, especially measured relative to the longest wing fringe setae (compare Figures 11C and 12C).  Figure 12A) with two small ventral teeth and a broad truncation dorsally. Maxillary palps two-segmented. Mid-lobe of mesoscutum ( Figure 12E) with fewer than 30 setae; each lateral lobe with three setae; each axilla with one seta; scutellum with four setae. Sculpture of mesoscutum aciculate; sculpture of axillae and scutellum longitudinal. Fore wing ( Figure 12C) with two setae on submarginal vein, 7-9 setae in basal cell, 7-11 setae on anterior margin of marginal vein, and one seta at the junction of the submarginal vein and parastigma. Linea calva absent. Submarginal 0.73x times marginal vein. Maximum length of fore wing 2.52x fore wing width, maximum width of fore wing 6.9x longest setae on marginal fringe. Ovipositor ( Figure 12F Remarks: Encarsia encantadora is morphologically closest to E. erwini, differing from that species mainly in having the third valvulae longer than the second valvifers. The fore wing is also broader in E. encantadora, especially measured relative to the longest wing fringe setae (compare Figures 11C and 12C).  Figure 13E) with about 18 setae; each lateral lobe with three setae; each axilla with one seta; scutellum with four setae. Sculpture of mesoscutum and axillae longitudinally aciculate; sculpture of scutellum longitudinal, transverse apically. Fore wing ( Figure 13C) with two large setae on submarginal vein and five smaller setae above, six setae in basal cell, seven setae on anterior margin of marginal vein, and one large seta at the junction of the submarginal vein and parastigma. Linea calva absent. Submarginal vein approximately equal in length to marginal vein. Maximum length of fore wing 2.9x fore wing width, maximum width of wing 3.75x longest seta on marginal fringe.
Distribution: ECUADOR: Napo. Remarks: Encarsia erwini appears to be most closely related to E. cylindrica, with which it shares the elongate antenna and lack of a linea calva. It differs from E. cylindrica in having far fewer setae on the mesoscutum. E. erwini is also morphologically close to E. encantadora, differing from that species mainly in having the third valvulae shorter than the second valvifers. The fore wing is also broader in E. encantadora, especially measured relative to the longest wing fringe setae.
Male. Color: Head light brown. Mesosoma and gaster uniformly brown but posterior third of mesoscutum, anterior third of axillae and scutellum yellow. Legs brown. Fore wings hyaline.
Morphology: Similar to that of female, except antennal formula, flagellum with longitudinal sensilla on all segments and funicle segments subequal in length. Maxillary palps two-segmented. Mid-lobe of mesoscutum ( Figure 16E) with 46-60 setae; each lateral lobe with two setae; each axilla with one seta; scutellum with four setae and two vestigial setal bases. Sculpture of mesoscutum aciculate; sculpture of axillae and scutellum longitudinal. Fore wing ( Figure 16C) with two large setae and 3-4 smaller setae on submarginal vein, 7-9 setae on anterior margin of marginal vein, and one seta at the junction of the submarginal vein and parastigma. Linea calva present. Submarginal equal to marginal vein; Maximum length of fore wing 2.54x fore wing width; maximum width of fore wing 6.7x longest setae on marginal fringe. Ovipositor (Figure 16F Remarks: Encarsia larensis appears morphologically closest to E. marynoyesae from which it differs in having a much longer funicle, and shorter ovipositor. No molecular data were available for this species. Chavez (1996)   Remarks: Encarsia marynoyesae is morphologically very similar to E. aisha in many respects (though distant to it based on DNA). The species can be distinguished by E. marynoyesae having the second valvifers almost 2x (1.8) the third valvulae; while they are 1.5x as long in E. aisha. In E. marynoyesae the clava is well over 2x the length of the funicle; in E. aisha it is less than 2x as long. E. marynoyesae also shares aspects of morphology with E. catula, but can be distinguished by having more than 30 setae on the mesoscutum, and V3 less than 1 2 the length of V2 (much more than 1 2 as long in E. catula). Sequence deposited under GenBank accession number: OQ683563.
Etymology. Named for Mary Noyes MBE (Member of the Order of the British Empire). 3. 3. 15. Encarsia mendesi (Polaszek and Hayat) comb. n. Female. Color: Antennae brown, paler on their ventral halves. Head dark brown with pale lines bordering the eyes and extending along the genae towards clypeus, antennal scrobes, a line from the apex of the scrobes to the median ocellus, and a transverse line midway between the antennal sockets and the median ocellus centrally bordering the dorsal end of antennal scrobes. Mesosoma and metasoma uniformly dark brown. Legs pale yellow, with coxae and hind femora dark brown. Wings infuscated below the submarginal and marginal vein; stigmal vein pale in contrast with a darker marginal vein.
Male. All aspects of coloration and morphology as for female, except the antennae and genitalic characters.
( Figure 22A-F) Female. Color: Antennae light brown slightly darker on F6 and radicle. Head brown with pale lines bordering the eyes and extending along the genae towards clypeus. Mesosoma dark brown but with scutellum and sides of the metanotum yellow. Metasoma light brown. Legs uniformly pale yellow. Wings infuscated below marginal vein, stigmal vein pale in contrast with a darker marginal vein.  Remarks: Encarsia svetlana is morphologically closest to E. venia from which it differs in having an F3 transverse, and its scape is entirely pale. Sequence data deposited under GenBank accession number: OQ683558.

Discussion
All taxa of Encarsia mexicana species-group recovered sister to Encarsia dichaeta have a linea calva present on the fore wing (absent in E. dichaeta; unknown D2672, and DNA.0165). The presence or absence of a linea clava may represent a local synapomorphy for these two clades within the mexicana-group. All taxa which lack a linea calva also lack the enlargement of the clava of the antenna common in this species-group, though Encarsia noora possesses a linea calva in the absence of an enlarged clava. Further phylogenetic analyses and sequence capture for the five other species without linea calva (E. diablejo, E. cylindrica, E. encantadora, E. erwini and E. napo) will be necessary to determine the informativeness of these characters. Other evident characters appear phylogenetically uninformative. Some surprises include E. svetlana appearing well-removed from E. venia while their morphology is extremely similar. E. aisha and E. noora despite being DNA sister species have markedly different flagellar shapes.
With the placement of Dirphys syn. n. in the middle of Encarsia, the genus expands to 473 described species. Establishment of the Encarsia mexicana species-group further increases the difficulty with which the genus can be recognized. In particular, the genus now contains species with a linea calva on the fore wing, and parasitoids with a gregarious life history, further expanding the already broad host range of the genus. The future classification of Encarsia at large will rely upon its clarification by robust phylogenetic hypotheses built upon large molecular datasets (Kresslein et al. unpublished). As the phylogeny of Encarsia is further resolved, it will be necessary to revisit the current classification of the genus and determine whether alternative generic classifications would allow for greater diagnosability of the taxa therein. Funding: This work was supported in part by grants from FUNDESIMCA Fundación para el Desarrollo e Impulso de las Ciencias Agrarias en Canaria.

Data Availability Statement:
All DNA sequence data that support the results and conclusions of this study can be found at GenBank.