PANGLAO 2004

PANGLAO 2004

Jengalpheops rufus gen. nov., sp. nov., is described on the basis of several specimens collected on intertidal sand and mud flats of Panglao I., the Philippines. The uniformly reddish shrimps inhabit burrow mounds of the large callianas-sid mudshrimp, Glypturus armatus (A. Milne Edwards, 1870). Jengalpheops is defined mainly by the dorsally exposed eyestalks; the absence of orbital teeth; the short triangular rostrum; the unspecialized and feebly enlarged chelipeds, with comb-like rows of setae on the carpus and without snapping mechanism on the fingers; the 3rd maxilliped with styliform lateral plate; and the presence of strap-like epipods on the 1st to 4th pereiopods. The new genus appears to be relatively basal within the family, and is presumably most closely related to Potamalpheops Powell, 1979 and Stenalpheops Miya, 1997.

PANGLAO 2004

Two new species of the alpheid shrimp genus Leptalpheus Williams, 1965, are described from the tropical western Pacific. Leptalpheus denticulatus, new species, is described on the basis of several specimens collected in Nhatrang Bay, Vietnam (type locality); Panglao, off Bohol, the Philippines; and Yaqara Bay, Viti Levu, Fiji. Leptalpheus dworschaki, new species, is described on the basis of two specimens collected in Panglao. All specimens were collected from burrows of callianassid ghost shrimps, including Glypturus cf. armatus (A. Milne-Edwards, 1870) and Neocallichirus calmani (Nobili, 1904). Both L. denticulatus, new species, and L. dworschaki, new species, are closely related to L pacificus Banner & Banner, 1974, from Hawaii, differing from the latter species in subtle morphological details.

PANGLAO 2004

Patagurus rex gen. et sp. nov., a deep-water pagurid hermit crab, is described and illustrated based on a single specimen dredged from 400 m off Moorea, Society Islands, French Polynesia. Patagurus is characterized by a subtriangular, vaulted, calcified carapace, with large, wing-like lateral processes, and is closely related to two other atypical pagurid genera, Porcellanopagurus Filhol, 1885 and Solitariopagurus Turkay, 1986. The broad, fully calcified carapace, calcified branchiostegites, as well as broad and rigidly articulated thoracic sternites make this remarkable animal one of the most crab-like hermit crabs. Patagurus rex carries small bivalve shells to protect its greatly reduced pleon. Carcinization pathways among asymmetrical hermit crabs and other anomurans are briefly reviewed and discussed.

TARASOC

Two uncommon, morphologically similar snapping shrimps, Alpheus percyi Coutiere, 1908 and A. pseudopugnax (Banner, 1953), are reported from various localities across the Indo-West and Central Pacific. Both species are re-diagnosed based on type specimens and recently collected material; a lectotype is designated for A. percyi. The range of A. pseudopugnax, previously known only from the type locality in Hawaii, is extended considerably to French Polynesia and Reunion Island in the western Indian Ocean. Both A. percyi and A. pseudopugnax present a significant variation in the postrostral region of the carapace, making identification of specimens based solely on morphology rather difficult. In the field, however, the two species can be easily separated by their diagnostic and very striking colour patterns. Colour patterns also suggest that A. percyi may be more closely related to the species of the A. gracilipes Stimpson, 1860 complex than to A. pseudopugnax.

PAPUA NIUGINI

The diagnostic characters of the laomediid mud shrimp genus Espeleonaushonia Juarrero & Martínez-Iglesias, 1997 are reassessed and the genus is synonymized with Naushonia Kingsley, 1897. A new species of Naushonia is described based on a single male collected at a depth of 16 m off the coast of Guadeloupe, French Antilles. Naushonia draconis sp. nov. is closest to two species of Naushonia formerly placed in Espeleonaushonia, namely Naushonia palauensis (Alvarez, Villalobos & Iliffe, 2010) comb. nov. and Naushonia augudrea (Juarrero & García, 1997) comb. nov., but can be separated from both of them and all other species of Naushonia by the high mid-dorsal carina, a row of very strong spines on the lateral surface of the carapace, and prominent mid-dorsal crests on the first, second and third pleomeres. A key to all known species of Naushonia is provided.

KARUBENTHOS 2012

Parabetaeus acanthus sp. nov. is described based on two specimens from Thuwal, Saudi Arabia, and Madang, Papua New Guinea. The new species is unique within the genus in possessing an anteriorly directed, spine-like tooth on the mid-dorsal line of the carapace, posterior to base of the eyes, and in the stylocerite not reaching the mid-length of the first article of the antennular peduncle. The chelipeds of P. acanthus sp. nov. are variable in size and proportions, as well as in the armature on the finger cutting edges. This cheliped polymorphism appears to be typical to all species of the genus.

PAPUA NIUGINI

MADEEP, PANGLAO 2004

The present study deals with four species of the alpheid shrimp genus Alpheopsis Coutière, 1897 characterised by the presence of at least one strong carina on the dorsal surface of the carapace. Alpheopsis trigona (Rathbun, 1901) is redescribed based on the holotype from Puerto Rico and additional material from US Virgin Islands and Florida. Two new species closely related to A. trigona, viz. A. paratrigona sp. nov. and A. gotrina sp. nov., are described, the first based on material from several localities in the tropical western Atlantic, and the second from the Pacific coast of Panama and Colombia. The three species together form a distinctive transisthmian clade within Alpheopsis, the A. trigona species complex, characterised by the presence of several strong longitudinal carinae on the carapace and very distinctive colour pattern. A more distantly related species, A. aristoteles sp. nov., characterised by the presence of only one strong mid-dorsal carina in the anterior region of the carapace, is described based on material from São Tomé Island in the tropical eastern Atlantic.

KARUBENTHOS 2012

Alpheus leptochiroides De Man, 1909, a poorly-known species originally described from the Kai Islands in eastern Indonesia, is reported from Kavieng, eastern Papua New Guinea, representing only the second record of this snapping shrimp and slightly extending its distribution range into the tropical western Pacific. The original description was based on a relatively young specimen, whereas the Kavieng specimen is clearly an adult male. Most importantly, several rather important characters of the species were omitted and/or not illustrated by De Man, including the unique and diagnostic rounded cuticular expansions on several areas of the third maxilliped, not present in any other alpheid shrimp. Therefore, a full redescription of the species is provided, with new detailed illustrations.

KAVIENG 2014

Three species of the alpheid shrimp genus Salmoneus Holthuis, 1955 associated with burrows of other decapod crustaceans are reported from various Indo-West Pacific localities. Salmoneus venustus sp. nov. is described based on material collected at two distant localities, Nha Trang Bay, southern Vietnam, the type locality of the new species, and the Yiti-Sifah region east of Muscat, northern Oman. Both specimens were collected with the aid of a suction pump applied to burrow entrances or mounds in muddy sand; the holotype was possibly associated with burrows of the callianassid ghost shrimp, Glypturus sp. Salmoneus venustus sp. nov. shares many characteristics with S. latirostris (Coutière, 1897), including the red banding of the pleon, but can be distinguished from S. latirostris and all other species of the genus by a unique combination of morphological characters. The large-sized Salmoneus brucei Komai, 2009 is reported from Sumba, central Indonesia, representing a significant southward extension of the species’ previously known distribution range and the first record since its original description. The callianassid ghost shrimp Lepidophthalmus cf. rosae (Nobili, 1904) is recorded as a new host of S. brucei. Finally, Salmoneus colinorum De Grave, 2004, associated with burrows of larger snapping shrimps from the Alpheus malabaricus Fabricius, 1798 species complex, is reported for the first time from Madang, Papua New Guinea, representing an eastward extension of the species’ previously known distribution range.

PAPUA NIUGINI

A new species of the alpheid shrimp genus Salmoneus Holthuis, 1955, probably an obligate associate of ghost shrimp burrows, is described based on material from Oman (type locality: Darsait near Muscat), Iran and the Philippines. Salmoneus durisi sp. nov. is characterised principally by both chelipeds enlarged, robust, with ventral and dorsal margins of chelae carrying long fine setae, and with minor chela fingers armed with a few large teeth on cutting edges. All specimens of Salmoneus durisi sp. nov. were collected either directly from burrows of larger decapod crustaceans with the aid of a suction pump, or by exposing burrows dug under large subtidal rocks. The Iranian specimen was found together with its presumed host, Neocallichirus calmani (Nobili, 1904). Two additional specimens from Indonesia and the Solomon Islands are tentatively assigned to S. cf. durisi sp. nov., awaiting further studies.

PANGLAO 2004

The alpheid shrimp Nennalpheus Banner & Banner, 1981, hitherto known only from a few records in the tropical western Pacific, is recorded for the first time from the tropical eastern Atlantic, as well as from the Indian Ocean. Nennalpheus gabonensis sp. nov. is described based on a single but complete male specimen collected in mediocre visibility conditions at 11 m depth on a rocky reef north of Libreville, Gabon. The new species presents a substantial number of morphological differences from the two other species of Nennalpheus, especially in the armature of the cheliped fingers. The previously known distribution range of Nennalpheus sibogae (De Man, 1910), the type species of the genus, is extended significantly to include the Central Pacific (Hawaiian Archipelago) and the Indian Ocean (Glorioso Islands in the Scattered Islands group, and Mayotte Island in the Comoro Archipelago). In addition, new illustrations of N. sibogae are provided, including the previously not figured mouthparts and high-quality, contrasting background colour photographs. Field observations indicate that Nennalpheus is able to produce a weak snapping sound by rapidly closing one or both chelipeds, a previously suspected, but never observed behaviour.

PAPUA NIUGINI

A new genus and species in the caridean shrimp family Palaemonidae is described based on three type specimens collected at a depth range of 208–385 m off Guadeloupe, French Antilles. Zoukaris festivus gen. et sp. nov. shares many characters with several western Atlantic deep-water species currently assigned to Periclimenes Costa, 1844, as well as with the monotypic western Atlantic genus Diapontonia Bruce, 1986 and the Indo-West Pacific genus Echinopericlimenes Marin & Chan, 2014. Zoukaris gen. nov. can be separated from all of them by a unique combination of morphological features, especially the configuration of the dactylus of the ambulatory pereiopods. In addition, Periclimenes milleri Bruce, 1986 is recorded from the French Antilles based on a single specimen, also from Guadeloupe; its colour pattern is illustrated for the first time.

KARUBENTHOS 2

Two new deep-water species of the snapping shrimp genus Alpheus Fabricius, 1798 are described based on the material collected by the expeditions BIOPAPUA, BOA1 and MIRIKY, organised by the Muséum National d’Histoire Naturelle in Paris. Alpheus alaincrosnieri n. sp. from the A. brevirostris (Olivier, 1811) group is described based on material dredged at depths of 198–408 m near the coasts of Papua New Guinea, Vanuatu and Madagascar. This species also occurs in the Philippines, based on morphological characters of a mutilated specimen from Masbate reported by Chace (1988). Alpheus alaincrosnieri n. sp. is unique within the A. brevirostris group, in having small orbital teeth. In most other features, A. alaincrosnieri n. sp. is morphologically closest to A. kagoshimanus Hayashi & Nagata, 2000, A. longipalma Komai & Ohtomi, 2018, A. macroskeles Alcock & Anderson, 1894, A. nonalter Kensley, 1969 and A. acutocarinatus De Man, 1909. Alpheus vanuatu n. sp. is described based on several specimens dredged at depths of 231–331 m off Espirito Santo, Vanuatu. This species most likely represents a deep-water member of the newly defined A. paracrinitus species group, sharing most morphological characters with A. tenuipes De Man, 1910 and A. labis Banner & Banner, 1982.

BIOPAPUA, BOA1, MIRIKY

A new alpheid shrimp genus, Crosnierocaris gen. nov., is established for Crosnierocaris athanasoides sp. nov., a very peculiar shrimp possibly associated with decomposing sea grass accumulations in deeper water. The type series of C. athanasoides sp. nov. was collected in the Mozambique Channel, north of Grande Glorieuse Island, at a depth of 240–255 m. The new genus presents a unique combination of morphological features, including the very long, slender rostrum armed with a subdistal ventral tooth; the pterygostomial angle projecting as a strong sharp tooth; the third to fifth pleura with their distoventral margins armed with two to four small teeth, a unique feature within the Alpheidae; the sixth pleuron with an articulated plate; the eyes largely exposed dorsally and laterally; the first pereiopods (chelipeds) feebly enlarged in both sexes, stouter but shorter than walking legs; the first pereiopod carpus with rows of short serrulate setae on its mesial surface; the second pereiopod carpus with five subdivisions; the third to fifth pereiopods with meri armed with stout cuspidate setae; the uropodal diaeresis with a strongly dentate lateral portion; and the complete gill-exopod formula, with mastigobranchs on coxae of the third maxilliped and first to fourth pereiopods. Within alpheid shrimps, the rostrum of C. athanasoides sp. nov. is one of the longest relative to the total body length, often reaching half of the carapace length. The phylogenetic position of Crosnierocaris gen. nov. within the Alpheidae remains to be determined, but based on its morphological characters, the new genus appears to be closest to Athanas Leach, 1814, especially in the configuration of the frontal area of the carapace, and Potamalpheops Powell, 1979, for instance, in the armature of the uropodal diaeresis and walking legs, as well as the presence of setal rows on the cheliped carpus. The new genus also shares several morphological features with Yagerocaris Kensley, 1988, which represents one of the least derived lineages within the family Alpheidae.

BIOMAGLO

Two new symbiotic species of decapod crustaceans, a palaemonid shrimp and a porcellanid crab, are described from the same type locality situated on the campus of the King Abdullah University of Science and Technology (KAUST) in Thuwal, on the Red Sea coast of Saudi Arabia. The palaemonid shrimp Palaemonella jamila sp. nov. is described based on one male and one female, whereas the porcellanid crab Enosteoides habibi sp. nov. is described based on three males and two females. Unfortunately, the originally collected and studied specimens were lost shortly after study. However, subsequent recollection of one specimen of P. jamila sp. nov. and one specimen of E. habibi sp. nov. enabled to deposit a holotype of each new species in the collections of the Florida Museum of Natural History, Gainesville, USA. Both species inhabit burrows of goby-associated snapping shrimps, Alpheus spp., in the shallow subtidal areas, at depths less than 1 m. In addition, a single female specimen of Palaemonella aliska Marin, 2008 collected at Al Fahal reef off KAUST represents the first record of the species in the Red Sea and the Indian Ocean, and a considerable range extension of its previously known distribution range. This species may be a close relative of P. jamila sp. nov., to which it is also ecologically similar. The present study highlights the necessity of prospecting the largely neglected and still undersampled shallow subtidal habitats of the Red Sea, and the Indo-West Pacific in general.

KOUMAC 2.1

The diversity and evolution of the Red Sea invertebrates in mesophotic and deep-water benthic ecosystems remain largely unexplored. The Palaemonidae is a diversified family of caridean shrimps with numerous taxa in need of taxonomic revisions based on recent molecular analyses. The Red Sea mesophotic and bathyal palaemonid shrimps are largely unstudied. During recent expeditions off the Red Sea coast of Saudi Arabia, several palaemonid specimens were collected at a depth range of 88–494 m, spanning the mesophotic and bathyal zones. This material was examined morphologically and genetically to infer phylogenetic relationships among the Red Sea taxa and several other palaemonid genera. The concordant morphological and genetic data led to the description of two new genera and two new species. Moreover, one species was recorded in the Red Sea for the first time, with a new host record, whereas three further deep-water species, which do not occur in the Red Sea, were formally transferred to a different genus. As more exploration efforts are deployed, research on the diversity and evolutionary relationships among marine invertebrates from the Red Sea will further underline the uniqueness of its mesophotic and bathyal fauna.

MUSORSTOM 1

Synalpheopsis gen. nov. is established for a remarkable new alpheid species, Synalpheopsis laureae sp. nov., presently known only from the male holotype collected at 111–162 m east of La Désirade, Guadeloupe, Lesser Antilles. Synalpheopsis gen. nov. peculiarly combines features of two genera, Alpheopsis Coutière, 1897 and Synalpheus Spence Bate, 1888; however, it is presumably closer to the former genus. The new genus is characterised by the moderately developed orbital hoods, well-developed rostrum and orbital teeth, sixth pleonite without articulated flap, tip of the third maxilliped with crown of spiniform setae, chelipeds with two strong teeth on distolateral margin and lacking snapping mechanism on finger cutting edges, and gill formula without mastigobranchs and setobranchs.

KARUBENTHOS 2

A new alpheid shrimp, Salmoneus ngae sp. nov., is described based on two specimens collected on a near-shore, seagrassrubble sandflat on the north-eastern coast of New Caledonia. The material from Lombok, Indonesia, previously reported as S. alpheophilus Anker & Marin, 2006, is reidentified as S. ngae sp. nov. The new species is a member of the informal S. gracilipes Miya, 1972 species group and appears to be a symbiont of spacious and complex burrows of the snapping shrimp Alpheus rapax Fabricius, 1798. Thus, S. ngae sp. nov. joins an ecological guild within the genus Salmoneus, which is composed of seven species associated with burrows of Alpheus spp. in various parts of the world and in different habitats.

KOUMAC 2.1, KOUMAC 2.2

The present study deals with several species of goby-associated snapping shrimp in the taxonomically challenging Alpheus brevirostris (Olivier, 1811) species group. Alpheus djeddensis Coutière, 1897 is redescribed based on two specimens from the original type series and material recently collected on the Red Sea coast of Saudi Arabia. Alpheus djiboutensis De Man, 1909, a species without extant type material, is removed from the synonymy of A. djeddensis after a careful analysis of the original description by De Man (1909). A cryptic species, morphologically undistinguishable from A. djeddensis, but with a significant divergence in the COI gene sequence and several consistent differences in colour pattern, is described as A. shukran sp. nov., based on material from Oman and Saudi Arabia. Both A. djeddensis and A. shukran sp. nov. are closely related to A. macellarius Chace, 1988, another potential species complex. Two species previously confused with A. djeddensis or A. djiboutensis in ecological literature and underwater field guides are described as A. tigrinus sp. nov. and A. berumeni sp. nov., based on material from the Red Sea, mainly Saudi Arabia. Each of them has a unique and diagnostic colour pattern and differs from A. djeddensis and A. djiboutensis by a few subtle morphological characters. The identity of the material preliminarily identified as A. cf. tigrinus and A. cf. berumeni from localities outside of the Red Sea needs to be confirmed by molecular analyses. Finally, three specimens from the Red Sea coast of Saudi Arabia are preliminarily identified as A. cf. djiboutensis; their true taxonomic identity, however, remains to be clarified further. All available information on goby associations for the herein treated species is provided.

KOUMAC 2.1, KOUMAC 2.3, PAPUA NIUGINI

In the present study, two deep-water species of the caridean shrimp family Alpheidae are described from two localities in the Indo-West Pacific, each based on a single specimen. Bannereus kebir sp. nov. is described based on a female holotype collected at a depth of 230‑243 m in the Kai Islands, Indonesia. The new species can be easily separated from the two congeners, B. anomalus Bruce, 1988 and B. chani Anker & Pachelle, 2020, by the presence of strap-like epipods on the first to third pereiopods. Salmoneus profundus sp. nov. is described based on an incomplete hermaphrodite holotype collected at a depth of 218‑225 m near Hanover Island, Papua New Guinea. This species differs from all other species of Salmoneus Holthuis, 1955 by the shape and armature of the enlarged minor cheliped, the elongate walking legs, and the absence of a strap-like epipod on the fourth pereiopod. It also represents by far the deepest-known member of Salmoneus, since all other known species in this genus occur at depths shallower than 90 m.

KARUBAR, KAVIENG 2014

The deep-water alpheid shrimp Batella praecipua De Grave, 2004 was originally described based on two incomplete specimens, both with their first chelipeds missing, collected at 400–440 m off New Caledonia. During the recent KANADEEP 1 expedition in the Coral Sea in 2017, the first complete specimen of B. praecipua was collected at a depth of 370–380 m. The morphology of this specimen is presented in detail, with emphasis on the highly diagnostic chelipeds, which are described and illustrated for the first time. A slight intraspecific variation of B. praecipua is reported and a DNA barcode (CO1) is provided for this species. The main differences between the three presently known species of Batella Holthuis, 1955 are discussed.

KANADEEP

The present study deals with the intricate phylogenetic relationships within an important clade of the caridean shrimp family Alpheidae, which includes the genera Salmoneus, Deioneus, and Triacanthoneus. Using molecular data from four genes (16S, 18S, COI, and H3), we herein provide the most comprehensive phylogenetic hypothesis for this alpheid clade, with 161 terminals representing 117 ESUs, i.e. genetically distinct species. Our analysis reveals four well-supported larger Clades, as well as 25 smaller clades, corresponding to species groups or species complexes. The results also show that the symbiotic lifestyle evolved multiple times within this clade, in some instances promoting convergent evolution. Ancestral character state analysis unveils extreme plasticity and independent shifts in the minor cheliped types. Furthermore, the impact of different lifestyles on the evolution of each minor cheliped type is discussed. Finally, we address the resulting paraphyly of Salmoneus and the intergeneric relationships between Salmoneus, Deioneus and Triacanthoneus.

KARUBENTHOS 2012, KAVIENG 2014, KOUMAC 2.3, MADIBENTHOS, PAPUA NIUGINI

Batellopsis paula gen. et sp. nov., a new hexactinellid-associated alpheid shrimp, is described based on a single specimen collected at a depth of 477–503 m north of Île des Pins, New Caledonia. Batellopsis gen. nov. is part of a clade of four genera all containing hexactinellid-associated species, for which a molecular phylogeny is presented. The evolution of several morphological characters, including orbital teeth, a bulge- fossa system on the fingers of the first pereiopod chela, and groups of microserrulate setae on the second pereiopod chela, is discussed in light of phylogenetic results.

EXBODI, KANACONO, KARUBAR, KOUMAC 2.1, MADIBENTHOS, TAIWAN 2002

KAVIENG 2014, PAPUA NIUGINI

KANACONO, KARUBAR, KOUMAC 2.1, PAPUA NIUGINI

PANGLAO 2004

PAPUA NIUGINI

PAPUA NIUGINI

The present study is the first major assessment of the marine decapod fauna of Ceará, northeastern Brazil, since contributions of J. Fausto-Filho in the 1960s–1970s. A fully updated checklist of all decapod crustaceans occurring in marine and estuarine habitats of Ceará is provided, based on literature records, specimens held in two carcinological collections of the Universidade Federal do Ceará (UFC), and material collected mainly by the authors between 2011 and 2014. A total of 337 decapod species are listed, distributed among the following taxa: Achelata (8 species), Anomura (42 species), Astacidea (1 species), Axiidea (11 species), Brachyura (162 species), Caridea (83 species), Dendrobranchiata (20 species), Gebiidea (9 species), and Stenopodidea (1 species). Among them, 23 species represent new records for Ceará, with 14 species, viz. Alpheus peasei (Armstrong, 1940), A. thomasi Hendrix & Gore, 1973, Ambidexter symmetricus Manning & Chace, 1971, Axianassa australis Rodrigues & Shimizu, 1992, Biffarius biformis (Biffar, 1971), B. fragilis (Biffar, 1970), Leptalpheus axianassae Dworschak & Coelho, 1999, L. forceps Williams, 1965, Lysmata bahia Rhyne & Lin, 2006, L. intermedia (Kingsley, 1878), Merhippolyte americana Holthuis, 1961, Neocallichirus maryae Karasawa, 2004, Ogyrides hayi Williams, 1981, and Typton carneus Holthuis, 1951, now having Ceará as the northern-most limit in their distribution range along the Brazilian coastline. One shrimp species, Lysmata lipkei Okuno & Fiedler, 2010, which was also found in the state of Rio Grande do Norte, possibly represents an invasive taxon in Brazil and the western Atlantic, originating from the Indo-West Pacific. Alpheus buckupi Almeida, Terossi, Araújo-Silva & Mantelatto, 2013, previously recorded from Ceará based on a colour photograph, is confirmed from this state, with specimens from several new localities. A few doubtful records from Ceará are briefly discussed. Colour photographs are provided for most of the taxa newly recorded from Ceará; some species are illustrated in colour for the first time.

KARUBENTHOS 2012

We use molecular analyses, color patterns, and records of distribution of mating pairs to reconstruct the global phylogeography of Alpheus lottini, a complex of cryptic coral-associated snapping shrimp species. Molecular data support the delineation of ancestral clades A, B, and C, and suggest five additional subdivisions within clades A and B. Clades A, B1, B2, and C exhibit color pattern differences and/or evidence of assortative mating, and thus merit species-level recognition. There is no evidence for assortative mating within clades A and B1, with likely reproductive compatibility (i.e., fertile clutches) in areas of sympatry. The clade diversity peaks in the Mariana Islands and the early branching clade C is restricted to the northern periphery of the Central and Western Pacific suggesting a Pacific origin of this group outside of the Coral Triangle. These findings underscore the prevalence of allopatric processes with possible ecological or microallopatric speciation in areas where clades overlap.

PAPUA NIUGINI