22 www-biodiversityjournal.com ISSN 2039-0394 (Print Edition) ISSN 2039-0408 (Online Edition) with the support of bo world biodiversity association o n I u Biodiversity Journal MARCH 2016, 7 (1): 1-200 FOR NATURALISTIC RESEARCH AND ENVIRONMENTAL STUDIES Polycera quadrilineata (O. F. Muller, 1776) - Eastern Sicily, Mediterranean Sea BIODIVERSITY JOURNAL 2016,7 (1): 1-200 Quaternly scientific journal edited by Edizioni Danaus, viaV. Di Marco 43,90143 Palermo, Italy www.biodiversityjournal.com biodiversityjournal@gmail.com Official authorization no. 40 (28.12.2010) ISSN 2039-0394 (Print Edition) ISSN 2039-0408 (Online Edition) EDITORIAL STAFF Managing Editor Ignazio Sparacio - Palermo, Italy Chief Editor Maria Stella Colomba University of Urbino “Carlo Bo”, Italy Secretary Fabio Liberto - Cefalu, Italy Marketing Editor Michele Bellavista - Palermo, Italy Assistant Editors David P. Cilia, Santa Venera, Malta Salvatore Giglio - Cefalu, Italy Armando Gregorini University of Urbino “Carlo Bo”, Italy Tommaso La Mantia - Univ. of Palermo, Italy Nunzia Oliva - Palermo, Italy Agatino Reitano - Catania, Italy Giorgio Sparacio - Palermo Fabio M.Viglianisi - University of Catania, Italy SCIENTIFIC COMMITTEE Vittorio Aliquo - Palermo, Italy Pietro Alicata - University of Catania, Italy Marco Arculeo - University of Palermo, Italy Paolo Audisio, Sapienza University of Rome, Italy Alberto Ballerio - Brescia, Italy Rostislav Bekchiev - National Museum of Natural History, Sofia, Bulgaria Christoph Buckle - Tubingen, Germany Attilio Carapezza - Palermo, Italy Donald S. Chandler - University of New Hampshire, Durham, U.S.A Renato Chemello - University of Palermo, Italy Giulio Cuccodoro -The Natural History Museum of Geneva, Switzerland Vera D'Urso - University of Catania, Italy Alan Deidun - University of Malta, Msida, Malta Gianniantonio Domina - University of Palermo, Italy Gerhard Falkner - Deutsche Malakozoologische Gesellschaft, Germany Paola Gianguzza - University of Palermo, Italy Ilia Gjonov -Sofia University, Bulgaria Adalgisa Guglielmino,Tuscia University, Viterbo, Italy Peter Hlavac - Prague, Czech Republic Ren Hirayama -Waseda University, Shinjuku-ku, Tokyo, Japan Rumyana Kostova - Sofia University, Bulgaria Sergey A. Kurbatov - Moscow, Russia Albena Lapeva-Gjonova - Sofia University, Bulgaria Oscar List - University of Catania, Italy Pietro Lo Cascio - Associazione “Nesos”, Lipari, Italy Nathalie Yonow - Swansea University, Swansea, Wales, U.K. Federico Marrone - University of Palermo, Italy Bruno Massa - University of Palermo, Italy Pietro Mazzola - University of Palermo, Italy David Mifsud - University of Malta, Msida, Malta Alessandro Minelli - University of Padova, Italy Pietro Minissale - University of Catania, Italy Marco Oliverio - University of Roma, Italy Roberto A. Pantaleoni - CNR National Research Council, Sassari, Italy Salvatore Pasta - Palermo, Italy Alfredo Petralia - University of Catania, Italy Roberto Poggi - Museo civico di Storia naturale“G. Doria”, Genova, Italy Francesco Maria Raimondo - University of Palermo, Italy Marcello Romano - Capaci, Italy Giorgio Sabella - University of Catania, Italy Danilo Scuderi - Catania, Italy Giuseppe FabrizioTurrisi - Catania, Italy Errol Vela - Universite Montpellier, France Polycera quadrilineata (O.F. Muller, 1776) (Gastropoda Polyceridae). Order Nudibranchia (Mollusca, Opisthobranchia). Nudibranchs are commonly known as “sea slugs” because they are not shelled molluscs. The evolution of the shell in gastropods followed a complexity plan of development, starting from simply low spiral, patelliform structures to highly twisted shells, the most safety house where a soft-body animal could hide from predators. How could shells be more efficient? After the “invention” of the shell, gastropods - which became heavy and slow - started to produce a thin shell. Increasing mobility conducted to shell reduction and this latter required a new plan of defense from predators. Probably around 3 or 4 hundreds of years ago, nudibranchs evolved from shelled molluscs and diversified. What is the successful of this new branch of gastropods due to? Toxicity or simply disgust to predators. This condition was reached by nudibranchs in two different ways. Some accumulate chemical active molecules throughout their tissues from the natural host upon which they feed, thus resulting venomous or stodgy. Some others build an internal equipment of spicules, which make them very hard to eat. How to infonn their potential predators of their dangerous internal items? Nudibranchs are very beautiful marine organisms, showing delicate external soft parts and spectacular colors, often comparable to butterflies. The reason of these showy colorations is the aposematic message; warning colorations mean: “I am venomous” so that predators immediately learn it is better to avoid these striking animals. The photograph shows a specimen of P. quadrilineata crawling on an ascidian looking for some encrusting bryozoans to eat (Summer 2004, Riposto, Catania, Eastern Sicily) (cover photo by Danilo Scuderi). Danilo Scuderi. Via Mauro de Mauro 1 5b, Belpasso, Catania; e-mail: danscu@tin.it Biodiversity Journal, 2016, 7 (1): 3-6 Nesting of the Black Stork Ciconia nigra Linnaeus, 1 758 (Aves Ciconiidae) in the FiumaraVitravo Valley (Calabria, Italy) Francesco Lamanna Alcimo street 88815 Strongoli Marina, Crotone, Italy; e-mail: niphargus@libero.it ABSTRACT The Fiumara Vitravo Valley in the province of Crotone in Italy, is a Site of National Interest for its rich biodiversity and peculiar habitat, and also a strategic area for the nesting of Black Stork, Ciconia nigra Linnaeus, 1758 (Aves Ciconiidae). The river morphology, the harshness of this wild territory, the luxuriant vegetation, the presence of a hydrographic network rich of trophic resources and the crucial position along the migratory routes, are fundamental for the reproductive biology and the evolution of this species. This work will expose the results of the monitoring activities that were carried out in 2015 by which it was possible to document the Black Stork nesting on rocky areas in the valley of Fiumara Vitravo. The ecological im- portance of the area is strongly in need of greater scientific attention and a suitable site pre- servation in order to favor the population increment of the Black Stork also in Calabria, where the active reproductive population was present only until 2001. The results are in evident countertrend with respect to older statistical data, which provide negative and sparse data for black stork presence in the “Alto Crotonese” region. KEY WORDS Ciconia nigra; Crotone; Calabria; nesting site. Received 03.12.2015; accepted 19.01.2016; printed 30.03.2016 INTRODUCTION The Black Stork, Ciconia nigra Linnaeus, 1758 (Aves Ciconiidae) is a bird with a wide territorial distribution. Its nesting area goes from Spain to Sachalin island between the 35° and 60° North parallel, with a separate population nesting in South Africa (Del Hoyo et al., 1992). The species, having a palearctic afro-tropical chorology, is very rare in western Europe, where it has suffered from drastic reduction with a complete disappearance in some states due to the destruction of its natural habitat. In Italy, the black stork is a migrating nesting species rarely wintering. Its biological characteristic is of long range flyer, able to travel over large portions of the sea, allowing it to migrate from win- tering zones to nesting areas travelling for thou- sands of kilometers. The populations move along not well defined routes, crossing the Mediterranean sea on a wide frontline. Some groups travel through the Strait of Gibraltar, others through the Red Sea along the Suez Canal to the Caucasian regions, others from the Black Sea go through the Bosphorus. One group crosses the eastern Mediterranean from Peloponnese partially exploiting the bridge fomied by the Egeo islands. A small group proceeds along the Sicily channel and the Italian peninsula (Petretti, 1993). 4 Francesco Lamanna The passage of Black Stork in Calabria is not well documented for the lack of an observation network throughout the territory. Small groups of isolated in- dividuals, observed during the passage, may lead one to think both the Tyrrhenian and Ionic side of Calabria as preferential migratory routes, although the crossing of the Sila plateau cannot be excluded. In general, the reproduction area of the species should include Eurasia, Southern Africa and Western Spain at the border with Portugal. Isolated populations are also found in central Europe and Balkans. The eastern reproduction area is more continuous including the north-east of Turkey, the Caucasus, and a wide region of Russia. In Italy, the first verified nesting was in 1994 in the natural park of Monte Fenera in the bassa Valsesia in the Pied- mont region. In the last years a gradual increment of the number of nesting couples has been observed in several Italian regions with a preference for the southern regions (Bordignon, 2006). THE BLACK STORK IN CALABRIA At the end of the 19th century Lucifero (2003), a man of wide cultural interests, published the first information on the presence of the black stork in Calabria. In that essay the Black Stork is classified as accidental and very rare and its presence was signaled in the area close to Crotone and Isola Capo Rizzuto. In the same essay some statements made by Moschella (in Lucifero, 2003), for the Reggio Calabria province, ensured the presence of the species in that region. The information was very scarce in the begin- ning of the century, and only starting from the 1970s, reliable data recorded the species as avail- able in the Calabria region. After 1970 the obser- vations became more frequent with several sightings. In 1994, the Black Stork nested in Ca- labria, with only one couple bringing four young birds to fly (Bordignon, 1995). The next year an- other couple brought two to flight. In 1996 no nesting was registered despite the presence of some individuals (G. Rocca personal observation) on the Lese and Neto rivers in the Crotone region. In 1997 only one couple was present bringing two young birds to flight. In the years 1998 and 1999 no nesting was registered but just the presence of isol- ated individuals on the Lese river (G. Rocca per- sonal observation). In 2000 only one couple nested in the Crotone region with four flying young birds (Bordignon et al., 2011). In 2001 the same nest was used by a couple for the deposition of four eggs and the flight of four young birds (Rocca, 2002). In the same year a second couple was detected by A. Digiorgio in the same reproduction area. In 2002 the presence of a couple with two immature individuals was registered in the nesting and feeding area. In March 2014, a serious event occurred in one of the most important migratory routes for migrating avifauna. In the core of the Parco Nazionale della Sila an adult black stork was found dead, shot by an unknown poacher in the S. Nicola location in the zone 2 of the park in the Serra Pedace district. In August 2014, during a research campaign, conducted by myself in the valley of Lese river, the presence of an isolated individual was detected. In February 2015, another disappointing event happened on the Amato river near Terzi di Lamezia Terme (Catanzaro) where one specimen was seen with a broken leg in an evident difficult condition but still able to fly (Lega Italiana Protezione Uccelli Sez. Rende, www.lipurende.it). In the present year an intensive search activity to individuate nesting black storks was successfully accomplished finding a couple regularly nesting on a rock face in the SIN of “Vallone Vitravo”. SIN (SITE OF NATIONAL INTEREST) “VALLONE VITRAVO” The Fiumana Vitravo is one of the major rivers of the “Alto Crotonese” district situated in the North East part of the Calabria region, having a major branch length of 43 Km. In its medium highest por- tion it has a torrential regime, while in the medium part water flows in a deep canyon. Downstream the morphology is like the Calabrian rivers’ with a wide bed and holm oaks. The site “Vallone Vitravo” (IT9300192), be- longing to the biogeographic mediterranean region with an abundance of wet fluvial habitat, includes 8 Km of riverbed of this important river extending in its median portion on a surface of about 800 ha. The area is characterized by a very dense ri- parian vegetation, with mixed forest of deciduous, sclerophyllous and brushwood, and Mediterranean Nesting of the Black Stork Ciconia nigra (Aves Ciconiidae) in the Fiumara Vitravo Valley (Calabria, Italy) 5 low. Ichthyic-fauna based on salmons populates the zones where water flows more rapidly and creates wide and deep potholes, while Cyprinidae stand in the valley areas. The biotic characterization of Vallone Vitravo was performed since the high naturalistic value of the site makes it a unique habitat for the preserva- tion of important floristic species, peculiar endemic floras and faunas and endangered birds. The geo- morphological characteristics of the area, with mighty and inaccessible rock walls, permit the nesting of animal species of the european com- munity interest included into the Attachment 1 of Direttiva “Uccelli” 79/409/CEE as Black Stork, Ciconia nigra , nesting area until 2001. THE NESTING SITE In August 2014, on the Lese river, close to the confluence with Neto river, a single individual of black stork was accidentally observed. It was an individual of which it was not possible to obtain any ethological information due to the late reproduction period and the difficulty in finding the feeding sites. In that circumstance the presence of any other indi- vidual or nesting site was not detected. This ap- pearance, of great ornithological importance, and related data on spring migration flows pushed us to plan for 201 5 a search campaign in the valley of the Fiumara Vitravo, nesting site of the species (Rocca, 2005). In May 2015 it was identified the pair and the nesting site. The nest was built within a natural cavity at the base of a shelf of rock, on a sandstone rock face in the valley of Fiumara Vitravo. The nest was at an altitude of 370 m on the see level at the top of the sandstone rock face which is 80 m long with East exposition. The great distance of the nest from the possible observation points, at least 300 m, together with the peculiar conformation of the valley, which barely offer a suitable observation prospective, did not permit to get information on the number of laid eggs. In the first decades of June, two nestlings, apparently one week old, were fed by both parents. In the last decade of July the feeding phase was regularly concluded and the young birds took their first flight. The nesting site were monitored visually from three observation points at a minimum distance of Figure 1. Black Stork flying over the Vitravo Valley. 300 m. After hatching, observations were made periodically with short cyclical 1 0 day visits on the sites in order to avoid disturbing the reproductive cycle of the couple. CONCLUSIONS N esting of black stork in the valley of Fiumara Vitravo brings the attention of the researchers to a site of greatest importance for the survival of this extraordinary bird. The reproduction success in the Alto Crotonese region shows, in this delicate phase of the geographic expansion of the species, a positive trend in the conquering of the habitats where black stork had disappeared for years. The natural preservation of these fragile and unique ecosystems imposes a collective effort to the scientific community. It should be necessary in the future to continue the monitoring of the site in order to remove or to reduce all the factors (pollu- tion, fire, anthropic impact, etc.) that limit the expansion of the species. REFERENCES Bordignon L., 1995. Prima nidificazione di cicogna nera, Ciconia nigra, in Italia. Rivista italiana di omitologia, 64: 106-116. 6 Francesco Lamanna Bordignon L., Branelli M., Buoninconti F., Caldarella M., Fraissinet M., Francione M., Fulco E., Gatti F., Marrese M., Rizzi V. & Visceglia M., 2011. La cicogna nera in Italia. Status e problemi di conser- vazione della popolazione nidificante, 2011. Del Hoyo J., Elliot A. & Sargatal J. (Eds.), 1992. Hand- book of the birds of the World. Vol. 1, Ostrichto Ducks. Lynx Edictions, Barcellona. Lucifero A., 2003. Avifauna e Mammiferi della Calabria, Selezione di Scritti Naturalistici. Greentime editori, Bologna, 60 pp. Petretti F., 1993. La nera sentinella dei boschi e delle rocce. Oasis, 9: 74-87. Rocca G., 2002. Nuovi dati sulla Cicogna nera, Ciconia nigra, in Calabria. Rivista italiana di ornitologia, 7 1 : 218-219. Rocca G., 2005. La cicogna nera in Calabria. In: Bor- dignon L. (Ed.). La cicogna nera in Italia. Parco Nat- urale del Monte Fenera. Tipolitografia di Borgosesia, Borgosesia (VC). Biodiversity Journal, 2016, 7 (1): 7-10 The amphioxus Epigonichthys maldivensis (Forster Cooper, 1 903) (Cephalochordata Branchiostomatidae) larvae in the plankton from Rapa Nui (Chile) and ecological implications Erika Meerhoff 1,2 *; David Veliz 2,3 ; Caren Vega-Retter 2,3 & BeatrizYannicelli 1,2 'Centro de Estudios Avanzados en Zonas Aridas (CEAZA), Coquimbo, Chile 2 Millennium Nucleus for Ecology and Sustainable Management of Oceanic Islands (ESMOI), Universidad Catolica del Norte, Lar- rondo 1281, Coquimbo, Chile 3 Departamento de Ciencias Ecologicas, Facultad de Ciencias, Universidad de Chile Corresponding author, e-mail: erikameerhoff@udec.cl ABSTRACT We report the first record of amphioxus larvae in the plankton from Rapa Nui island (Chile). Zooplankton was sampled using an oblique Bongo net during an oceanograhic survey in April and September 2015. A total of four larvae were collected in the coastal area of Rapa Nui in April and 13 in September. The larvae were identified as Epigonichthys maldivensis (Forster Cooper, 1903) (Cephalochordata Branchiostomatidae) using both morphological and genetic characters. The water column in this area presented a mean temperature of 21.2°C, a mean salinity of 35.7 %o and 4.94 ml/L dissolved oxygen in April, and 20°C and 35.75 %o mean salinity in September. Amphioxus have been reported as playing a key role in marine food webs transferring important amounts of microbial production to higher trophic levels, due to this their role in the Rapa Nui plankton and benthos as adults could be interesting because Easter island is located in the oligotrophic gyre of the South Pacific ocean where a microbial trophic web is expected to dominate. This record increases the biodiversity of Rapa Nui plankton and widens the geographic distribution of E. maldivensis that was restricted only to the Western and Central Pacific and Indian Ocean. KEY WORDS amphioxus larvae; Pacific Ocean; plankton. Received 03.12.2015; accepted 19.01.2016; printed 30.03.2016 INTRODUCTION The Amphioxus or lancelets (Chordata) com- prise the subphylum Cephalochordata (Schubert et al., 2006); which is formed by three genera: Bran- chiostoma Costa, 1 834, Epigonichthys Peters, 1876 and Asymmetron Andrews, 1893 (Konetal., 2007). The amphioxus are filter-feeding marine organisms that as adults burrow in the sand, gravel or shell deposits in tropical and/or temperate waters around the world ocean (Bertrand & Escriva, 2011). The filtering is performed through jawless ciliated mouths (Vergara et al., 2011). Amphioxus are found in general in shallow wa- ters close to the shore (0.5 to 40 m depths) and many species prefer habitats of coarse sand and gravel (Desdevises et al., 2011). They live in a vari- ety of coastal habitats, estuaries, coastal lagoons, open coasts and river deltas (Laudien et al., 2007; Chen, 2008). However, little is known about the ecological role of these organisms (Vergara et al., 201 1). In addition, some amphioxus have been con- sidered as endangered species (Kubokawa et al., 1998). Environmental factors as temperature and salinity changes are determinant in the life cycle of some amphioxus species (Webb, 1956a; Webb, 8 Erika Meerhoff etalii 1956b; Webb & Hill, 1958). As a consequence, the amphioxus populations migrate between winter and summer (Webb, 1971), and the larvae are described as restricted to waters of high salinity and temper- ature (Webb & Hill, 1958). The duration and timing of the spawning season varies between species (Stokes & Holland, 1996; Holland, 2011). When the gametes are released in the water, fecundation occurs and the embryos persist in the plankton (Ber- trand & Escriva, 2011) until metamorphosis, when they migrate to the sand and become benthic adults. Some authors have studied the zooplankton and meroplankton around Easter Island (Castro & Landaeta, 2002; Palma & Siva, 2006; Mujica, 2006) most zooplankton results are from CIMAR islands cruise in November 1999. However, there are no records of amphioxus larvae or adults in the area. In this work we describe the presence of amphioxus larvae from Rapa Nui plankton (Chile) for the first time. Larvae were found in stations close to the coast around the island in April and September 2015. MATERIAL AND METHODS Zooplankton samples and hydrographic meas- urements were gathered in the coastal area of Easter Island or Rapa Nui (27° 13’ S - 109°37’ W), Chile, in April and September 2015. The hydrographic characterization of the water column was done using a set of CTD profiles in both months (Seabird 1 8). Zooplankton samples were collected by oblique tows from a depth of 300 m up to the surface, using a Bongo net with 300 pm mesh and 60 cm mouth diameter. The volume of sampled water was estim- ated using a mechanical flowmeter (General Ocean- ics) attached to the net. Samples were preserved in 96% ethanol, until laboratory identification and quantification. In these samples seventeen amphi- oxus larvae were found. Considering that no in- formation about amphioxus larvae morphology is available, three larvae were used to perform the genetic identification. After that a simple morpho- logical description of the larvae is also supplied. Genetic identification Three larvae were used for the genetic analysis. The DN A extraction was conducted using the Qiagen QIAamp kit (Mississauga, Canada). The mitochon- drial COI gene was amplified using the protocol and primers described by Folmer et al. (1994) with 56°C as annealing temperature. Forward and reverse sequencing was performed at Pontificia Universidad Catolica de Chile and aligned by eye using the ProSeq v.2.9 software (Filatov, 2002). The haplotype was deposited in Genbank (Accession Number: KU201542). The Blast tool was used to determine similarities with sequences deposited in Genbank. In order to determine the nucleotide relationship among lancelets, a neighbour-joining based phylo- genetic (NJ) analysis was performed using Mega 6.0 software (Tamura et al., 2013). Using a boot- strap of 10,000 replicates, the analysis tested the consistency of each branch in the tree, grouping sequences with similar nucleotide composition. Using this method, unidentified sequence obtained in this study could be grouped with conspecific sampled in other geographical areas. RESULTS AND DISCUSSION A total of 4 amphioxus larvae were found in the coastal area of Rapa Nui in April and 13 in Septem- ber 2015. The larvae were identified as Epi- gonichthys maldivensis (Forster Cooper, 1903) (Cephalochordata Branchiostomatidae) (Fig. 1). In April, in the south station, larvae were found up to 200 m depth and the abundance was 0.8 individual per 1000 m 3 , while in the south-east station, the abundance of E. maldivensis larvae was 2 indi- viduals per 1000 m 3 and were found between 300 m depth and surface. The amphioxus larvae mean abundance in September was 2 individuals per 1000 m 3 and they were found in the south station. The environmental characteristics of the area were mean water temperature of 21.2°C, mean salinity of 35.7 %o and 4.94 ml/L dissolved oxygen in April, and 20°C and 35.75 %o mean salinity in September. Genetic identification. One haplotype of 550 bp was obtained for the larvae. The analysis of the COI gene showed a clear relationship of our sequence with Epigonichthys maldivensis (Fig. 2). The Blast ana- lysis showed a similarity of 99% with one sequence of E. maldivensis (Accession Number: AB1 10093. 1), deposited by Nohara et al. (2005) and obtained from one individual collected in the Kuroshira Island, Japan. Both sequences differ only in 6 bp. Epigonichthys maldivensis is a tropical species whose distribution was restricted only to the Western and Central Pacific and Indian Ocean The first record of the amphioxus Epigonichthys maldivensis larvae in the plankton from Rapa Nui (Chile) 9 (Richardson & McKenzie, 1994; Poss & Boschung, 1996; Lin et al., 2015), the present results expand the geographic range of this species to Rapa Nui island. Lancelets exhibit a week- to month long planktonic larval stage (Wiclcstead, 1970; Wu et al., 1994; Stokes & Holland, 1996) and in Eastern Island these were present in April and September 2015. The benthic communities from Rapa Nui are extremely species-poor compared with reefs in the central and western Pacific (Friedlander et al., 2013), the presence of the amphioxus larvae, implies that amphioxus adults probably live in the benthos that would contribute to the benthos species richness. Moreover, anecdotal histories from the local fisherman of Rapa Nui reporting, in some areas and dates, the presence of white filaments like hairs in the bottom, are likely to corroborate our findings; these filaments could be the adult amphi- oxus. This record increases the biodiversity value of Rapa Nui. In addition, since amphioxus have been reported as playing a key role in marine food webs transferring important amounts of microbial production to higher trophic levels (Chen et al., 2008), their role in the Rapa Nui plankton and benthos as adults could be interesting since Easter island is located in the oligotrophic gyre of the South Pacific ocean where a microbial trophic web is ex- pected to dominate. Finally, new amphioxus genome sequences will be of great importance for compar- ative genomics at the inter and intra species levels. ACKNOWLEDGEMENTS Authors acknowledge the support from the Chilean army at Easter Island and the ORC A diving center to conduct the samplings. EM acknowledges the support from Postdoctoral-FONDECYT/Chile 3150419. EM acknowledges the support of Millen- nium Nucleus for Ecology and Sustainable Man- agement of Oceanic Islands (ESMOI). CV acknowledges the support of Fondecyt de Iniciacion N° 11150213. Authors also acknowledge Carolina Paz Concha Molina from CFRD University of Concepcion for her contribution with the drawing. Figure 1. Above: schematic views of the amphioxus larva, basic anatomy the oral cirri, the segmented muscles, and the notochord are signaled. Below: Epigonichthys maldivensis larval individual collected from Rapa Nui. 49 18 17 38 39 E? 10 100 93 | Epigonichthys maldivensis 1 He I Epigonichthys maldivensis2 Ernaldrvensis Epigonichthys cultellusl 100 1 Epigonichthys cultellus2 -Branchiostoma betchen'2 - Branchiostoma beichen3 1Q0 100 3 □□ | — Branchiostoma malayanum2 I Branchiostoma maiayanum3 Branchiostoma floridae3 ^ | — Branchiostoma fioridael S5^ — Branchiostoma florrdae2 — Branchiostoma japomcum3 p Branchiostoma japonicuml 70 L Branchiostoma japomcum2 Branchiostoma lanceolatum3 100 i Branchiostoma lanceolatuml 100 < Branchiostoma lanceolatum2 100 lAsymmetron spl ' Asymmetron sp2 ICO lAsymmetron mfeaiml I Asymmetron inferum2 ( ■ Asymmetron lucayanuml Asymmetron Iuc3yanum2 Asymmetron Iucayanum3 Figure 2. Neighbour-joining tree of the COI sequences for the Branchiostomidae species. The number at the tree nodes indicates the bootstrap values from 10,000 replicates. The figure shows also the GenBank Accession Numbers. REFERENCES Bertrand S. & Escriva H., 2011. Evolutionary crossroads in developmental biology: amphioxus. Development, 138: 4819-4830. Castro L.R. & Landaeta M.F., 2002. Patrones de dis- tribucion y acumulacion larval en tomo de las islas oceanicas: Islas de Pascua y Salas y Gomez. Ciencia y Tecnologia del Mar, 25: 131-145. 10 Erika Meerhoff etalii Chen Y., Shin P.K.S. & Cheung S.G., 2008. Growth, secondary production and gonad development of two co-existing amphioxus species ( Branchiostoma belcheri and B. malayanum ) in subtropical Hong Kong. Journal of Experimental Marine Biology and Ecology, 357: 64-74. Desdevises Y., Maillet V., Fuentes M. & Escriva H., 20 1 1 . A snapshot of the population structure of Bran- chiostoma lanceolatum in the Racou Beach, France, during its spawning season. PLoS ONE 6, el 8520. Folmer S.C., Black M., Hoek R., Lutz R.A. & Vrijenhoek R. , 1994. 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Biodiversity Journal, 2016, 7 (1): 11-16 Description of three new subspecies of Carabus Linnaeus, 1 758 (subgenus Coptolabrus Solier, 1 848) and taxonomic changing on some Carabus from Far East of Russia (Coleoptera Cara- bidae Carabinae) Ivan Rapuzzi Via Cialla 47, 33040 Prepotto, Udine, Italy; email: info@ronchidicialla.it ABSTRACT Three new Carabus Linnaeus, 1758 (subgenus Coptolabrus Solier, 1848) subspecies from Far East of Russia and Central China (Anhui Province, Chongqing Province) are described and figured: C. ( Coptolabrus ) smaragdinus losevi n. ssp., C. ( Coptolabrus ) elysii wangguofeni n. ssp. and C. ( Coptolabrus ) ignigena tenuitarsatus n. ssp. Comparative notes with the closest taxa are provided. Carabus ( Morphocarabus ) hummeli vladobydovi Obydov, 2007, C. (A ulonocarabus) gossarei mareschii Rapuzzi, 2010, C. ( Megodontus ) vietinghoffii rugicolor Rapuzzi, 2010 and C. ( Coptolabrus ) smaragdinus robinzoni Rapuzzi, 2010 recently con- sidered as synonyms are resurrected as valid subspecies. KEY WORDS Carabus', Coptolabrus; new subspecies; Far East Russia; China; taxonomic changing. Received 22.01.2016; accepted 24.02.2016; printed 30.03.2016 INTRODUCTION The study of some Coleoptera Carabidae of the genus Carabus Linnaeus, 1758 (subgenus Copto- labrus Solier, 1848 see Hauser, 1921, 1932a, 1932b; Deuve & Font, 1998; Deuve, 2004) pre- served in the author's collection in part provided by Mr. Oleg Losev (Pavlovo, Russia) and Mr. Xi Huangshun (Shanghai, China) gives the opportunity to individuate three new subspecies: C. ( Copto- labrus ) smaragdinus losevi n. ssp. from South Primorye in the Far East of Russia, C. ( Copto- labrus ) elysii wangguofeni n. ssp. from Anhui province, Central China and and C. ( Coptolabrus ) ignigena tenuitarsatus n. ssp. from Chongqing province, Central China. In the second part of this paper five Carabus taxa recentely considered as synonyms by Sun- dukov (2013) are resurrected as valid subspecies. RESULTS New taxa Carabus ( Coptolabrus ) smaragdinus losevi n. ssp. Examined material. Holotype: 1 male, Far East of Russia, South Primorye, Khasanskiy district, Furugelm Island, 11/13.VII.2013, O. Losev legit; preserved in the author’s collection. Paratypes: 6 males and 3 females, Far East of Russia, South Primorye, Khasanskiy district, Furugelm Island, 11/13.VIL2013, 0. Losev legit; 6 males and 3 females, Far East of Russia, South Primorye, Khasanskiy district, Krabbe peninsula, 30.VI/11.VII.2012, O. Losev legit; 25 males and 3 females, Far East of Russia, South Primorye, Khasanskiy district, Krabbe peninsula, 7/18.VII.2013, O. Losev legit; 24 males and 3 females, Far East of Russia, South- 12 Ivan Rapuzzi west Primorskiy region, Khasanskiy district, Mramomyy cape env., 42°34’N; 130°48’E, 28.VII/12.VIIL2012, A. Plutenko legit. The paratypes are preserved in the author’s col- lection, O. Losev collection and A. Plutenko collec- tion (Russia). Description of Holotype male. Length includ- ing mandibles: 31 mm, maximum width of elytra: 9.8 mm (Pig. 1). Head and pronotum cupper-red, elytra cupper-red with cupper-green sides, relatively shiny; primary and secondary relieved intervals of elytra black. Ventral side of pronotum and epipleura cupper-red, metallic, abdomen dark violet; palpi antennae and legs black. Head elongate; surface strongly and uniformly punctured; supra-antennary ridge bent upwards; clypeus relieved, lateral ridges very deep and punctured. Mandibles very long and thin, of “cychrisanf ’ shape. Eyes emispheric and prominent. Labrum bilobate, multi-setulose. Very long and developed palpi, sub-apical segment of labial palpi bi-setose; apical segment of maxillary and labial palpi dilated. Antennae thin, extending with 4 antennomers beyond the base of pronotum and extending more or less the third of elytra. Disc of pronotum nearly flat; sides of pronotum narrow margined, slightly bent upwards at the base; hind angles rounded and very slightly protruding behind its base; surface of pronotum uniformly and very densely punctured, faintly roughly. Elytra quite elongate, oval, very convex, maximum width at the middle; shoulders narrow, slightly pronounced; sculpture triploid heterodyname type: primary intervals forming tubercles of oval shape, smooth; secondary smaller, rounded and veiy smooth; tertiary completely reduced. Legs very long and strong. Aedeagus: the median lobe in lateral view (Fig. 2) is regularly curved, apex long and curved; dorsal view in figure 3. Variability. Paratypes have a small variability: the length of the body ranges from 27.5 mm to 32 mm for the males and from 27 mm to 34 mm for the females. The colour of the specimens from Krabbe peninsula is cupper green; the specimens from Furugelm island and Mramornyy cape have constantly the holotype colour. Etimology. This new interesting Coptolabrus subspecies is very cordially dedicated to Mr. Oleg Losev (Pavlovo, Nizhegorodskaya region, Russia) who collected part of the specimens. Remarks. The small size, the convex shape of elytra with smooth intervals, the quite transverse and of hexagonal shape pronotum, the very small and elongate head and the dominant cupper-red colour characterize the new subspecies. From C. ( Coptolabrus ) smaragdinus mandschu- ricus Semenov, 1898 the new subspecies is distin- guish for the smaller size and for the sculpture of elytra formed by larger and smoother intervals. From C. smaragdinus coreicus Hauser, 1921 the new subspecies is geographically separate by the large Tumen Jiang river valley and differs for the smaller head, the smaller size, the longer mucrons of elytra, the larger pronotum and smoother sculp- ture of elytra. The closest subspecies is C. smaragdinus robin- zoni Rapuzzi, 2010 described from Reyneke Island near Vladivostok (Rapuzzi, 2010; 2012). With the new subspecies it shares the same small size but differs for the dominant red colour, the transverse pronotum of hexagonal shape, the very convex elytra, the less raised sculpture of elytra and for the shape of aedeagus more curved with longer apex. Carabus ( Coptolabrus ) elysii wangguofeni Ra- puzzi et Huangshun n. ssp. Examined material. Holotype: male, China, Anhui province, Taihu, Wangling vill., South slope of Mt. Dabieshan, 400 m, 10/30.IV.2015, (30°31T8" N; 116°16'39" E), Xihuangshun legit; preserved in Ivan Rapuzzi collection. Paratypes: 9 males and 11 females, China, Anhui province, Taihu, Wangling vill., North slope of Mt. Dabieshan, 400 m, 10/30.IV.2015, Xihuangshun legit; 3 females, idem, except V.2014; the paratypes are preserved in Ivan Rapuzzi collection. Description of Holotype male. Length in- cluding mandibles: 41 mm, maximum width of elytra: 13 mm (Fig. 4). Upper surface metallic, dull; head green; pronotum and side of elytra gold- green; disc of elytra olive green; primary and secondary intervals of elytra black. Ventral side of pronotum and epipleura green, metallic, abdomen dark violet; appendix black. Head elongate; surface strongly punctured, frons convex and punctured; clypeus very sparsely punctured; clypeus fovea deep and punctured. Mandible long, sickled shape. Palps long with the apical segment strongly dilated; Three newsubspecies of Carabus (Coptolabrus) and taxonomic changing on some Carabus from Far East of Russia 13 Figures 1-3. Carabus ( Coptolabrus ) smaragdinus losevi n. ssp. holotype male. Fig. 1: holotype. Fig. 2: holotype male aedeagus: median lobe in lateral view. Fig. 3: idem, apex in dorsal view. Figures 4-6. Carabus ( Coptolabrus ) elysii wangguofeni n. ssp. holotype male. Fig. 4: holotype. Fig. 5: holotype male aedeagus: median lobe in lateral view. Fig. 6: idem, apex in dorsal view. penultimate segment of labial palps bi-setose. Pronotum of hexagonal shape, transverse (1.21 times as long as broad); base of pronotum large; sides quite rounded, marginated, bent upwards; basal lobes large and rounded, protruding its base; surface of pronotum densely and shallow punc- tured. Elytra oval; disc convex; mucrones short; sculpture triploid heterodyname type: primary tubercles rounded and close; secondary smaller and rounded; tertiary forming grains strongly rough; ground roughly sculptured. Legs quite short. Male aedeagus (Figs. 5,6). Variability. Very variable in colour: green, bluish-green, blue, golden-green; the margins often differ from the discs of pronotum and elytra; colour of head and pronotum often contrasting with that of elytra. The colour always has cold tints. The length of the body ranges from 37 mm to 41 mm for the males and from 40 mm to 44 mm for the females. One female specimens has the sculpture of elytra with tubercles more elongate. Etimology. The beautiful new Coptolabrus taxa is very cordially dedicated to Mrs. Wang Guo- fen (Shanghai, China) wife of Mr. Xi Huangshun. The co-author of this new subspecies is Huangshun Xi from Shanghai, China Remarks. From Southern Anhui several Copto- labrus taxa are known: Carabus ( Coptolabrus ) elysii elysii Thomson, 1846: Ngang-Wei, Anking (= Anhui, Anqing) (Hauser, 1921); Carabus ( Coptolabrus ) elysii connectens Hauser, 1912: Ngang-Wei, sudlicher Teil (= Anhui, Southern part) (Hauser, 1921); Carabus ( Coptolabrus ) elysii anhweiensis Hauser, 1932: Anking (= Anqing) (Hauser, 1932a loc. typ.; 1932b). Very close to C. elysii connectens it is considered as a synonym by Brezina (2003); Carabus ( Coptolabrus ) lafossei tungchengensis Li, 1993: Tongcheng Xian, Longming, Shanling, locus typicus (Li, 1993); Carabus ( Coptolabrus ) lafossei dabieshanus Imura, 1996: Anhui: Dabie Shan, Yuexi, Mt. Miaodaoshan, locus typicus (Imura, 1996); Hetupu (Deuve, 1997); Qianshan Xian, Tianzhu Mt. (Imura, 1996); Qian Shan; Jiuhua Shan; Baima Jian 14 Ivan Rapuzzi (Kleinfeld, 1997). Very close to C. ( Coptolabrus ) lafossei tungchengensis it is considered as a syn- onym by Brezina (2003) Carabus ( Coptolabrus ) lafossei jingdensis Deuve et Li, 2006: Anhui, Jingde Xian, Junle, 30°20’N; 118°30’E, locus typicus (Deuve et Li, 2006). From the adjacent area were described: Carabus ( Coptolabrus ) lafossei tiantai Klein- feld, 1997: NE-Hubei: Hong’an, Mt.Tiantai, 31:23N/114:37E, locus typicus (Kleinfeld, 1997); Carabus ( Coptolabrus ) lafossei pseudocoelestis Kleinfeld, 1999: N-Hubei, Shuizhou, Dahong Mt., 3 1 :29N/1 12:58E, 1200 m, locus typicus (Kleinfeld, 1999); Carabus ( Coptolabrus ) elysii pulcher Kleinfeld, 1997: S-Henan, S of Xinyang, Jigong Shan, 31:49N/114:06E, locus typicus (Kleinfeld, 1997). The closest form is C. elysii pulcher from which the new subspecies is easily distinguished by the following characters: smaller size, very different colour with domination of cold tints; larger pro- notum with smoother sides (less angulate); more convex elytra; shorter elytral mucrones; primary intervals forming smaller and nearly perfect rounded tubercles. From C. elysii elysii and C. elysii anhweiensis the new subspecies differs by: larger size; more elongate and slender body shape; hexagonal pro- notum; rounded and raised primary tubercles (smoother in C. elysii elysii and C. elysii anhweien- sis); longer elytral mucrones. The range of the new subspecies is geograph- ically very close to that of C. lafossei dabiesanus but very easily distinguishable by several strong characters: different colour (in C. lafossei dabies- anus constantly with black elytra and dark blue elytra margins, head and pronotum); more trans- verse and less angulate pronotum; upper surface of head and pronotum strongly punctured (smooth in C. lafossei lafossei ); different sculpture of elytra and shorter mucrons of elytra. From C. lafossei tiantai, C. lafossei pseudoce- lestis and C. lafossei jingdensis the new taxon has all the distinctive characters of the species that permit to separate C. elysii elysii from C. lafossei lafossei. Carabus lafossei tiantai and the new sub- species show, in part, the same colour. Carabus ( Coptolabrus ) ignigena tenuitarsatus n. ssp. Examined material. Holotype: male, China, Chongqing province, Pengshui county, Mt. Heimending, local collector legit; preserved in the author’s collection. Paratype: 1 male, China, Chongqing province, Pengshui county, Mt. Heimending, local collector legit; the paratype is preserved in the author’s collection. Description of holotype male. Small size and very thin shape for the species, length includ- ing mandibles 38.5 mm; maximum width of elytra 11.8 mm (Fig. 7). Upper surface metallic, rather mat; head with supra antennary ridges green; pro- notum with sides gold green, disk darker; elytra uniformly green, sides very shine, brilliant; primary and secondary intervals black. Ventral face of head black; ventral face of pronotum and epipleura dark green, metallic; abdomen black with violet shades, metallic; appendix black. Head long and very slender; surface of head densely punctured, frons very convex. Mandibles elong- ate. Eyes quite small and slightly salient. Palpi long with the apical segment strongly dilated; penultimate segment of labial palpi bisetose. Pro- notum long and very narrow for the species, as broad as long, sides of pronotum very sinuate, rounded; hind angles salient and very few pro- truding behind the base; upper surface flat; surface of pronotum densely punctured, median sulcus very superficial. Elytra narrow and very elongate for the species, ovals; disc convex. Primary intervals perfectly rounded or slightly elongate, very prominent; secondary forming aligned grains; tertiary reduced. Long mucrones. Legs quite short. First and second protarsal segments of male slightly dilated with complete adhesive soles; the third male protarsal segment not dilated and with very rudimental adhesive soles. Male aedeagus (Figs. 8, 9) is characteristic for the species but quite slender and of narrower shape. Variability. No significant variability of the paratypes Etimology. The new subspecies is named after the slightly dilated male protarsal segments. Three newsubspecies of Carabus (Coptolabrus) and taxonomic changing on some Carabus from Far East of Russia 15 Remarks. As expected the new taxa is morpho- logically close to the northern most subspecies of C. ignigena : C. ( Coptolabrus ) ignigena cristiano- fonti Deuve et Font, 2008 and C. ( Coptolabrus ) ignigena tongrenensis Deuve et Li, 2006. From C. (C.) ignigena cristianofonti, that it is the closest form, it is easily distinguished by the following characters: slender shape of head and pronotum; sides of pronotum sinuate but not angled; much elongate elytra with primary intervals more relieved; protarsal segments of male slightly dilated, the third segment not dilated and with very rudimental adhesive soles. From C. (C.) ignigena tongrenensis the new subspecies is distinguished by the following characters: smaller size; slender shape of head and pronotum; primary intervals of elytral sculpture more prominent; shorter legs; protarsal segments of male slightly dilated, the third segment not dilated and with veiy rudimental adhesive soles. Up to now the new subspecies is the northernmost population of the whole range of C. ignigena and it is the first record of the species for the Chongqing province. Figures 7-9. Carabus ( Coptolabrus ) ignigena tenuitarsatus n. ssp. holotype male. Fig. 7: holotype. Fig. 8: holotype male aedeagus: median lobe in lateral view. Fig. 9: idem, apex in dorsal view. Taxonomic notes Recently Sundukov (2013) established as syn- onyms four Carabus subspecies described from the Peter the Great Gulf Islands, Vladivostok area, Far East of Russia: C. (Morph ocarabus) hummeli smaragdulus Kraatz, 1878 = C. (M.) hummeli vladobydovi Obydov, 2007); Carabus (Aulono- carabus ) gossarei gossarei Haury, 1879 = C. (A.) gossarei mareschii Rapuzzi, 2010; Carabus (Megodontus) vietinghoff bowringi Chaudoir, 1 863 = C. (M.) vietinghoff rugicolor Rapuzzi, 2010 and C. ( Coptolabrus ) smaragdinus mandschuricus Semenov, 1898 = C. (C.) smaragdinus robinzoni Rapuzzi, 2010. For the significant morphological characters and the perfect isolation under insular conditions all these taxa will be resurrect: - Carabus (Morphocarabus) hummeli vladoby- dovi Obydov, 2007 stat. resurr. Described from Popov Island (Obydov, 2007) C. hummeli vladoby- dovi has good morphological characters that permit to separate it from the populations from the main- land as well as from C. hummeli putyatini Rapuzzi (2012) from Putyatin island. Carabus hummeli vladobydovi differs from all the other known hum- meli subspecies for its veiy peculiar coloration: violet-pink or red-pink pronotum, pink with gold or green shades elytra and purple margins. - Carabus (Aulonocarabus) gossarei mareschii Rapuzzi, 2010 stat. resurr. Described and known only from the Askol’d Island C. gossarei mareschii is easily separable from C. gossarei gossarei by several characters: larger size and more developed elytra of ovate-elongate shape. The pronotum is less punctate with larger and dipper basal impressions. Elytral sculpture with less interrupted and less prominent primary intervals. Male aedeagus longer and larger with the median lobe more developed. - Carabus (Megodontus) vietinghoffii rugicolor Rapuzzi, 2010 stat. resurr. Described from Reyneke Island it is one of the most distinctive subspecies of C. vietinghoffii. It is easily distinguished from C. vietinghoffii bowringi by significant and constant characters: in general bigger and stronger shape; very different colour: upper surface dark red to black-violet, rather mat, margins of elytra of the same colour. Male aedeagus differs for: in lateral view the median lobe is more developed and the apical lobe is longer; apex in frontal view curved on the left. 16 Ivan Rapuzzi - Carabus ( Coptolabrus ) smaragdinus robinzoni Rapuzzi, 2010 stat. resurr. Described from Reyneke Island it differs from C. smaragdinus mandschuri- cus by the following characters: smaller size; slender and flatter shape; pronotum as broad as long, not transverse; stronger elytral sculpture; apical lobe of male aedeagus longer and slender. It is interesting to note that C. smaragdinus robinzoni is very constant in his type locality. REFERENCES Brezina B., 2003. World Catalogue of the Genus Carabus L. Pensoft, Sofia-Moscow 1999, 170 pp. Deuve Th., 1997. Catalogue des Carabini et Cy- chrini de Chine. Memories de la Societe ento- mologique de France, 1: 1-236. Deuve Th., 2004. Illustrated Catalogue of the Genus Carabus of the World (Coleoptera, Carabidae). Pensoft. Sofia-Moscow, 461 pp. Deuve Th. & Font M.L., 1998. Descriptions de deux noveaux Coptolabrus du Guizhou (Cole- optera, Carabidae). Coleopteres, 4: 59-63. Hauser G., 1921. Die Damaster-Coptolabrus- Gruppe der Gattung Carabus. Zoologische Jahrbiicher. Abteilung fur Szstematik, 45: 1- 394. Hauser G., 1932a. Neue Coptolabrus- Formen. Mitteilungen der deutschen entomologischen Gesellschaft, 3: 8-9 Hauser G., 1932b. Weitere Mitteilungen zur Kenntnis der Coptolabrus- Formen. Deutsche entomologische Zeitschrift, 76: 96. Imura Y., 1996. Contribution to the knowledge of carabid fauna (Coleoptera, Carabidae) in West- ern Hubei and Southwestern Anhui, China. Geklcan-Mushi, 229: 10-14. Kleinfeld F., 1997. Zwei neue Carabus ( Copto - labrus)- Unterarten aus dem Dabie-Shan, China (Provinzen Hubei, Hunan) (Coleoptera: Cara- bidae: Carabini). Entomologische Zeitschrift, 107: 459-464. Kleinfeld F., 1999. Neue Taxa der Gattung Carabus 0 Coptolabrus , Apotomopterus, Pagocarabus, Rhigocarabus ) aus den chinesischen Provinzen Henan, Hubei und Sichuan (Coleoptera: Cara- bidae: Carabini). Entomologische Zeitschrift, 109: 1-16. Li J.K., 1993. Carabidae. In: Studies on fauna and ecogeography of soil animal ( Coptolabrus : 138— 139). Obydov D., 2007. Two new subspecies of Carabus {Morpho carabus) hummeli Fisher von Wald- heim, 1823 from Ussuri Land (Coleoptera, Carabidae). Lambillionea, 107: 153-158. Rapuzzi I., 2010. Descrizione di tre nuove sotto- specie di Carabus provenienti dalle isole al largo di Vladivostok nell’Estremo Oriente russo (Coleoptera, Carabidae). Lambillionea, 110: 310-314. Rapuzzi I., 2012. Preliminary notice on the genus Carabus Linnaeus, 1758 (Coleoptera, Cara- bidae) of the islands of Peter the Great Gulf in the Far East of Russia, Primorski province, Vladivostok area with description of a new sub- species. Biodiversity Journal, 3: 479-486. Sundukov Yu.N., 2013. An annotated catalogue of the ground beetles (Coleoptera: Caraboidea) of Sikhote-Alin. Vladivostok: Dalnauka, 271 pp. Biodiversity Journal, 2016, 7 (1): 17-20 On the presence of the Andaman lobster, Metanephrops andamanicus (Wood-Mason, 1891) (Crustacea Astacidea Nephropidae) in Palabuhanratu bay (S-Java, Indonesia) Yusli Wardiatno 1 *, AgusAlim Hakim 1 , Ali Mashar 1 , Nurlisa Alias Butet 1 , LukyAdrianto 1 &Achmad Farajallah 2 'Department of Aquatic Resources Management, Faculty of Fisheries and Marine Science, Bogor Agricultural University, Kampus IPB Darmaga, Bogor 16680, West Java, Indonesia. department of Biology, Faculty of Mathematics and Natural Sciences, Bogor Agricultural University, Kampus IPB Darmaga, Bogor 16680, West Java, Indonesia. *Corresponding author: email: yusli@ipb.ac.id ABSTRACT The first Andaman lobster, Metanephrops andamanicus (Wood-Mason, 1891) (Crustacea Astacidea Nephropidae) record from south of Java waters, part of Indian Ocean is reported in this paper. A total of 3 specimens were collected at a fish harbor in Palabuhanratu bay in May 2015. Morphological characters are illustrated and described. This finding enhances the biodiversity lists of Indonesian crustaceans. KEY WORDS Andaman lobster; Decapoda; Indian Ocean; Java Island; morphological descriptions. Received 23.01.2016; accepted 02.03.2016; printed 30.03.2016 INTRODUCTION The lobsters of the family Nephropidae are deep-sea forms and commonly found at depths from 150 to more than 1893 m (Chang et al., 2014). In general nephropid lobsters are bottom-dwellers with a preference for soft sediments, and living within their self-made burrows is the biological behavior in some species (Chan, 1998). The family Nephropidae currently includes 57 species belonging to 14 genera (Holthuis, 1991; Chan, 1998; Tiirkay, 2001; Chan, 2010; Ahyong et al., 2012; Chan et al., 2014). Previously, genus Metanephrops Jenkins, 1972 was divided into four morphological groups, namely thomsoni (Bate, 1888), binghami (Boone, 1927), arafurensis (De Man, 1905) and japonicus (Tapparone-Canefri, 1873) (Holthuis, 1991). However, with molecular analysis approach, Chan et al. (2009) refuted mono- phyly of the arafurensis and thomsoni groups. Among the groups, japonicus has the highest num- ber of species. Some of the current researches on Indonesian crustaceans, reported the presence of first records species, especially hippoid crabs, such as Albunea symmysta (Linnaeus, 1758) (Mashar et al., 2015), Hippa marmorata Hombron et Jacquinot, 1846) (Wardiatno et al., 2015), Hippa adactyla Fabricius, 1787 (Ardika et al., 2015). This paper presents a new record of the Anda- man lobster, Metanephrops andamanicus (Wood- Mason, 1891) from south of Java, Indonesia. MATERIAL AND METHODS Three M. andamanicus specimens were collec- ted in May 2015, from a fish harbor in Palabuhan- ratu bay, District Sukabumi, South of Java, Indonesia (Fig. 1). They were preserved in 96% 18 Yusli Wardiatno etalii alcohol and taken to the laboratory for analysis. Identification was based on the morphological characters using taxonomic key books from FAO (Holthuis, 1991; Chan, 1998). One example of the specimens is presented in figure 2. The specimens were lodged in the Department of Aquatic Re- sources Management, Bogor Agricultural Univer- sity, Indonesia. RESULTS SYSTEMATICS Infraorder ASTACIDEA Scho Its et Richter, 1995 Family NEPHROPIDAE Dana, 1852 Genus Metanephrops Jenkins, 1972 Metanephrops andamanicus (Wood-Mason, 1891) Examined material. 3 males: carapace length 51.04, 55.97, and 57.20 mm, total length 141.82, 149.34, and 154.23 mm, weight 65, 78, and 88 gram. 17.V.2015, Palabuhanratu fishing harbor, South of Java, Indonesia. Diagnosis. Carapace ofM andamanicus smooth between ridges and large spines (Fig. 3). Eyes large and black, postrostral carinae with three teeth (Fig. 4). Surface of abdominal tergites conspicuously sculptured; raised parts of dorsal surface of abdominal somites smooth and naked; second to fifth abdominal somites with marked dorsomedian carina, flanked by pair of conspicuous longitudinal grooves (Fig. 5). Fifth abdominal somite without distinct spines on carina separating tergite from pleuron. Dorsomedian carina of sixth abdominal somite without submedian spines. Spine in middle of lateral margin of sixth abdominal somite short, tip far from posterolateral margin of somite. Chelae of first pereiopods heavily ridged and spinulose, without large spines; no prominent basal spine on outer edge of movable finger of large chela. Inner margin of merus of first pereiopod weakly spinu- lose (Fig. 6). Distribution. Indo-West Pacific region: East Africa (Tanzania, Zanzibar, Kenya and Somalia), the Andaman Sea, the South China Sea (not includ- ing the Philippines), and Indonesia, and perhaps also Papua New Guinea (Holthuis, 1991; Chan, 1998; Tshudy et al., 2007). DISCUSSION Holthuis (1991), Chan (1998) and Tshudy et al. (2007) revealed the distribution of M. andamanicus in Indo-West Pacific region from eastern Africa to the Andaman Sea, the South China Sea (but not the Philippines), Indonesia, and perhaps also Papua New Guinea. According to the IUCN Red List of Threathened Species the occurence of the species in Indonesia was reported in Kalimantan, Sumatra and Sulawesi. However, in a short survey on May 2015 we could find this species in Palabuhanratu bay located in south of Java and it is a new record. Some lobster species were previously reported from several parts of Indonesia, and they were highly valuable species, such as Panulirus penicillatus (Olivier, 1791) (Chow et al., 2011; Kalih, 2012; Abdullah et al., 2014), Linuparus somniosus Berry et George, 1972 (Wowor, 1999), P. versicolor (Latreille, 1804) (Ongkers et al., 2014), P. homarus, (Linnaeus, 1758), P. longipes (A. Milne-Edwards, 1868), P ornatus (Fabricius, 1798), Parribacus antarcticus (Lund, 1793) (Kalih, 2012). Con- sequently, the presence of M. andamanicus in Palabuhanratu bay increases the list of lobster biod- iversity in Indonesian waters. In fishery point of view, some species of genus Metanephrops have commercial potential and become the deep water fishery targets lobster and catched by trawl; those species are M. mozambicus (Macpherson, 1990) in Africa (Fennessy & Groeneveld, 1997; Groeneveld & Everett, 2015), M. thomsoni in northern part of the East China Sea (Choi et al., 2008), M. challengeri (Balls, 1914) in New Zeland (Tuck et al., 2015), M. andamanicus in east coast of Southern Africa (Mutagyera, 1979). In the fish market located in Palabuhanratu bay, south of Java M. andamanicus can be regularly found indicating its economical value in the area. As fishery target, biological information of this species is needed for its sustainable management. Exploration in biological aspects of M. andamani- cus is open for future studies. AKNOWLEDGEMENTS The research was funded by Indonesian Gover- nment through Directorate General of Higher Edu- cation, Ministry of Education and Culture from Andaman lobster, Metanephrops andamanicus (Crustacea Nephropidae) in Palabuhanratu bay (S-Java, Indonesia) 19 Figure 1. Map of Java Island with the insert map of Indonesia. Palabuhanratu bay is indicated by open-square and pointed with an arrow. Figure 2. Metanephrops andamanicus (male) collected from a fish harbor in Palabuhanratu Bay, south of Java, Indonesia. Figures 3-6. Metanephrops andamanicus , south of Java (Indian Ocean), male (carapace length 55.97 mm). Fig. 3: carapace, lateral view. Fig. 4: carapace, dorsal view. Fig. 5: abdomen, dorsal view. Fig. 6: first pereiopod. Scale bars 10 mm. 20 Yusli Wardiatno etalii Fiscal Year 2015. The authors wish to thank to Mr. Agus for his assistance during specimen collection. REFERENCES Abdullah M.F., Alimuddin, Muththalib M., Salama A.J. & Imai H., 2014. Genetic isolation among the North- western, Southwestern and Central-Eastern Indian Ocean populations of the pronghorn spiny lobster Panulirus penicillatus . International Journal of Molecular Sciences, 15: 9242-9254. Ardika P.U., Farajallah A. & Wardiatno Y., 2015. First record of Hippa adactyla (Fabricius, 1787; Crusta- cea, Anomura, Hippidae) from Indonesian Waters. Tropical Life Sciences Research, 26: 105-110. Ahyong S.T., Webber R. W. & Chan T.Y., 2012. Thymops takedai, a new species of deepwater lobster from the Southwest Atlantic Ocean with additional records of ‘thymopine’ lobsters (Decapoda, Nephropidae). In: Komatsu H., Okuno J. & Fukuoka K. (Eds.), Studies on Eumalacostraca: a homage to Masatsune Takeda. Series Crustaceana Monographs. Brill, 49-61. Chan T.Y., 1998. Lobster. In: Carpenter K.E. & Niem V.H. (Eds.), FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians and sharks. Rome, FAO, 687-1396. Chan T.Y., 2010. Annotated checklist of the world’s marine lobsters (Crustacea: Decapoda: Astacidea, Glypheidea, Achelata, Polychelida). The Raffles Bulletin of Zoology, 23: 153-181. Chan T.Y., Ho K.C., Li C.P. & Chu K.H., 2009. Origin and diversification of the clawed lobster genus Metanephrops (Crustacea: Decapoda: Nephropidae). Molecular Phylogenetics and Evolution, 50: 411-422. Chang S., Chan T.Y. & Ahyong S.T., 2014. Two new species of the rare lobster genus Thaumastocheles Wood-Mason, 1874 (Reptantia: Nephropidae) dis- covered from recent deep-sea expeditions in the Indo- West Pacific. Journal of Crustacean Biology, 34: 107-122. Choi J.H., Kim J.N., Kim M.H., Chang D.S., Yoo J.T. & Kim J.K., 2008. Population biology and feeding habits of the nephropid lobster Metanephrops thom- soni (Bate, 1888) in the East China Sea. Journal of Environmental Biology, 29: 453-456. Chow S., Jeffs A., Miyake Y., Konishi K., Okazaki M., Suzuki N., Abdullah M.F., Imai H., Wakabayasi T. & Sakai M., 2011. Genetic isolation between the Western and Eastern Pacific populations of pronghorn spiny lobster Panulirus penicillatus. PLoS ONE, 6: e29280. Fennessy S.T. & Groeneveld J.C., 1997. A review of the offshore trawl fishery for crustaceans on the east coast of South Africa. Fisheries management and ecology, 4: 135-147. Groeneveld J.C. & Everett B.I., 2015. Deep water- water trawl fisheries for crustaceans. In: van der Elst R.P. & Everett B.I. (Eds.), Offshore fisheries of the South- west Indian Ocean: their status and the impact on vul- nerable species. Oceanographic Research Institute and Western Indian Ocean Marine Science Associ- ation. Special Publication No. 10. Chapter 3, 68-119. Holthuis L.B., 1991. FAO species catalogue. Vol 13. Marine lobsters of the world. An annotated and illus- trated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis. 125. Vol. 13. Rome. FAO, 292 pp. Kalih L.A.T.T.W.S., 2012. Diversity and distribution of Palinurid and Scyllarid lobster in the Lombok island coastal waters. Thesis. Graduate School, Gadjah Mada University, Yogyakarta. Mashar A., Wardiatno Y., Boer M., Butet N.A., Farajallah A. & Ardika P.U., 2015. First record of Albunea symmysta (Crustacea: Decapoda: Albuneidae) from Sumatra and Java, Indonesia. AACL, 8: 611-615. Mutagyera W.B., 1979. On Themis orientalis and Meta- nephrops andamanicus (Macrura, Scyllaridae and Nephropidae) off Kenya coast. East African Agricul- tural and Forestry Journal, 45: 142-145. Ongkers O.T.S., Pattiasina B.J., Tetelepta J.M.S., Natan Y. & Pattikawa J.A., 2014. Some biological aspects of painted spiny lobster (Panulirus versicolor ) in Latuhalat waters, Ambon Island, Indonesia. AACL, 7: 469-474. Tshudy D., Chan T.Y. & Sorhannus U., 2007. Morpho- logy based cladistic analysis of Metanephrops'. the extant genus of clawed lobster (Nephropidae). Journal of Crustacean Biology, 27: 463M76. Tuck I.D., Parsons D.M., Hartill B.W. & Chiswell S.M., 2015. Scampi (Metanephrops challenged ) emergence patterns and catchability. ICES Journal of Marine Science, 72 (Suppl. 1): i 1 99 — i2 1 0 . Tiirkay M., 2001. Decapoda. In: Costello M.J., Emblow C. & White R.J. (Eds.), European register of marine species: a check-list of the marine species in Europe and a bibliography of guides to their identification. Collection Patrimoines Naturels, 50. Museum na- tional d’Histoire naturelle: Paris, 284-292. Wardiatno Y., Ardika P.U., Farajallah A., Mashar A. & Ismail, 2015. The mole crab Hippa marmorata (Hombron et Jacquinot, 1846) (Crustacea Anomura Hippidae): a first record from Indonesian waters. Biodiversity Journal, 6: 517-520. Wowor D., 1999. The spear lobster, Linuparus somniosus Berry & George, 1972 (Decapoda, Palinuridae) in Indonesia. Crustaceana, 72: 673-684. Biodiversity Journal, 2016, 7 (1): 21-24 Does local knowledge change after a species long term ab- sence? The case of giant river otters Pteronum brasiliensis Gmelin, 1 788 (Carnivora Mustelidae) O. Eric Rami'rez-Bravo Departamento de Ciencias Qiumico-Biologicas, Universidad de las Americas, Puebla, Santa Catarina Martir, SinNumero, Cholula, Puebla; e-mail: ermex02@yahoo.com ABSTRACT Public participation could be useful to determine species presence and ecological aspects, however it is possible that local knowledge of species whose populations had suffered a decrease could have changed. To determine current knowledge of giant river otter, Pteronura brasiliensis Gmelin, 1788 (Carnivora Mustelidae), we undertook a preliminary assessment based on 35 interviews preformed between June and August 2014 with natural resources users in the Pacaya-Samiria Reserve (Peru) aimed to determine the presence, feeding habits, re- production periods, and threats. It was possible to determine that current knowledge cor- respond with available information in literature thus, I consider that it is possible to use public participation in cases of little known species that are recovering. KEY WORDS Public monitoring; Pteronura', Pacaya-Samiria Reserve; endangered species. Received 03.01.2016; accepted 31.01.2016; printed 30.03.2016 Habitat loss, fragmentation, and degradation, along with other human-related causes have put most ecosystems and the species that inhabit them at risk (Myers, 1988). Therefore, conserva- tion strategies rely on the prioritization of areas that are key for the long term survival of many species. Such prioritization becomes more important in areas with high biodiversity; and even more so when these areas are related with high human dens- ities, where actions are needed sooner rather than later (Sanderson et al., 2002). However, information on both ecosystems and species at regional level is often missing making it necessary to generate a strategy that could help to increase knowledge at this level. It has been proposed that this kind of information can be obtained from the general public as a first step for management; as scientific research is usually limited in space and time (i.e. short term studies in a specific site) making some changes to go either unperceived or identified after a huge gap of time (Scholte, 2011). Just to mention, in Mada- gascar local knowledge has been used to shape distribution of carnivore species (Kotschwar et al., 2015) and in Zimbabwe to determine population trends of different carnivore and game species (Gandiwa, 2012). However, there is not enough information on how local knowledge and perception changes after a charismatic species disappears from a region such as in the case of the giant river otter, Pteronura brasiliensis Gmelin, 1788 (Carnivora Mustelidae). This species was once distributed in most fresh- water streams of South America, from Venezuela to Argentina (Eisenberg, 1989). Its numbers have 22 O. Eric Ramirez-Bravo decreased significantly up to the extent that some populations have disappeared from its former range due hunting and habitat loss (Carter & Rosas, 1997; Recharte & Bodmer, 2010). The species is currently listed as endangered by the red list (IUCN, 2016) with a projected population decrease of about 50% within the next twenty years (Shostell & Ruiz- Garcia, 2013). Fortunately, due a ban on hunting and a decrease on its commercial demand in Peru, giant river otter populations have increased in certain areas such as in the Yavari River (Recharte & Bodmer, 2010) and in the Pacaya-Samiria Na- tional Reserve in Pern (Groenendijk et al., 2001). The species is important at local level as it is con- sidered a top predator and due its potential as a bio indicator as it is especially sensitive to disturbance and resource availability, preferring conserved areas with good fish stocks (Carter & Rosas, 1997; Groen- endijk et al., 2001; Recharte & Bodmer, 2010). Un- fortunately, scientific information of the species along its range is scarce except for a few areas (e.g. Madre de Dios: Flajek & Groenendijk, 2006) and Pacaya-Samiria National Reserve (Groenendijk et al., 2001). However, there is not enough informa- tion on how local knowledge and perception changed after the long term absence of the species in the region. Thus, it is important to determine if users of natural resources are aware of the giant Figure 1. Pacaya-Samiria National Reserve in Northeastern Peru. river otter presence and ecology in order to include their knowledge in management plans. I undertook semi structured interviews with the natural resource users of the Pacaya-Samiria Na- tional Reserve in northeastern Peru to assess their actual knowledge on giant river otters (Fig. 1). The Pacaya-Samiria National Reserve is located in the Amazon Basin and is considered the largest protec- ted area of flooded forest in the Amazon with 20,800 km 2 (Bodmer et al., 2011). Its average annual rainfall is 2000-3000 mm and a mean tem- perature between 20 and 33°C (Bodmer et al., 2011). The reserve and its buffer zone have 203 rural settlements; most of them (89%) are small villages with less than 500 inhabitants located on the borders of the Maranon and Ucayali/Puinahua rivers (Gonzalez, 2003). The households in the area include people of mixed origins (mestizos), natives from the ethnic groups Cocama-Cocamilla and Shipibo-Conibo, whose major economic activities include fishing, agriculture, game hunting, and extraction of forest products (Gonzalez, 2003). Some communities have been actively involved in groups of natural resources management in the Reserve (Puertas et al., 2000; Piana et al., 2003). I concentrated my efforts in the Samiria River, a black water river preferred by giant river otters (Carter & Rosas, 1997). I used the vigilance point 2 known as “Tacshacocha” as interviewing place, since visitors and members of the community man- agement groups have to register when travelling upriver. I made a total of 35 interviews between June and August 2014. The survey consisted in a set of 22 questions aimed to determine the presence of the species, habitat preferences, reproduction patterns and potential threats to giant river otters. Inter- viewed persons belong to five different communit- ies: Leoncio Pradro (47%), San Martin de Tipishca (29%), San Carlos (12%), Santa Rita (3%) and Victoria (9%). On average, the interviewees were 41 years old. Half of them (50%) belong to one of the local community-based conservation groups which were formed aiming for the sustainable use of natural resources as well as turtle management and conservation; they also serve as guides for scientific groups (19%). Sixty-nine percent of the interviewees typically use the reserve throughout the year, another 22% use it only during the dry season. Therefore, I considered that responses were based on field experience. Does local knowledge change after a species long term absence? The case of giant river otters Pteronura brasiliensis 23 Locals stated that otters can be observed throughout the year (40% of interviewees), but that are easier to detect during the dry season (43%). These observations can be explained because their movements are concentrated to lakes and rivers during the dry season, while they move to flooded forests and small creeks during the wet season (Hajek & Groenendijk, 2006), becoming scattered and harder to detect. Moreover, 53% of the respond- ents identified both river and rainforest as preferred habitats and another 41% considered river as the main one. Locals also reported diurnal observations (54% of interviewees), especially during the early hours of the morning (34%). This is supported by Carter & Rosas (1997), who identified giant river otters as diurnal. Interviewees reported that the main activities conducted by the river otters were playing, feeding, fishing, and vigilance, which correspond with previous reports about their daily activity (Carter & Rosas, 1997; Hajek & Groenendijk, 2006). The diet of the giant river otter varies with habitat type and species diversity in the area (Hajek & Groenendijk, 2006). Fish are the main diet component (Carter & Rosas, 1997; Hajek & Groenendijk, 2006), but other groups such as mam- mals and crabs have also been recorded (Hajek & Groenendijk, 2006). Preferred fish consumed by the giant river otter belong to the suborders Characoidei (characins), Percoidei (perch) and Siluroidei (cat- fish) (Carter & Rosas, 1997). Accordingly, inter- viewees identified fish of different species such as carachama ( Pseudorinelepis spp.) and different types of piranha (Characoidei) as the main diet component of the species. Knowledge about reproduction and cub devel- opment tended to vary. All interviewees considered that otters reproduce in the Samiria River, but only 80% have seen cubs. Eighty-eight percent of locals reported that otters breed during the dry season (May-September), which corresponds with obser- vations in other areas (Duplaix, 1980; Hajek & Groenendijk, 2006). Although litter size is known to vaiy between one and five cubs per season (Carter & Rosas, 1997; Hajek & Groenendijk, 2006), locals have little knowledge about this fact as just 29% consider that the species had just one cub per year. It is noteworthy that interviewees claimed they cannot differentiate pregnant from non-pregnant females (62%); but they can differen- tiate adults and cubs by size (76%). The success on conservation measures that have resulted in river otter population increase (Recharte & Bodmer, 2010) has been noted by interviewees, where 91% considered that the otter population was growing. Otter population increases may lead to a raise in human-otter conflicts. In fact, 49% of them considered the species as harmful to fishing nets and fish stocks, 46% stated that locals are afraid of the giant river otter, and 21% reported known previous attacks to humans. This corresponds with observations made by Carter & Rosas (1997), where people in recently colonized areas of the Amazon forest feared giant river otters. Inter- viewees suggest fears result from lack of know- ledge about the species. Regardless, 91% of the interviewees claimed local communities know that the giant otter is protected. Also, 91% of the re- spondents considered the giant otter as an important and emblematic species in the area because it represents the reserve and is part of the ecosystem. This is supported by the fact that 92% of them reported that this species is no longer hunted in the area as it is extremely prohibited, except occasion- ally when cubs are captured to be kept as pets or to be sold to zoos, as previously reported (Duplaix, 1980; Carter & Rosas, 1997). Our results indicate that, despite the species is still at low densities and was carried almost to the point of local extinction, local people who visit this reserve are well informed about the presence, eco- logy and distribution of the species. The latter can be confirmed by comparing published information from zoos and field observations about the species with local knowledge. In the specific case of the giant river otter, results showed that it is possible to use local knowledge as baseline information to generate conservation projects and community projects. Thus, it can be considered that for regions with limited information about species and eco- systems it is possible to use public participation, especially of community conservation groups, where available, in order to generate management plans or even monitoring programs. ACKNOWLEDGEMENTS I would like to thank the Servicio Nacional de Areas Naturales Protegidas (SERNANP) and to Fundamazonia for all their support during the field work. 24 O. Eric Ramirez-Bravo REFERENCES Bodmer R., Puertas P., Antunez M., Fang T. & Gil G., 2011. Impacts of climate change on wildlife conservation in the Samiria River Basin of the Pacaya-Samiria National Reserve, Peru, http:// opwall.com/research-library/research-reports/ peru/Accessed on October 12, 2014 Carter S.K. & Rosas F.C.W., 1997. Biology and conserva- tion of the giant otter Pteronura brasiliensis . Mammal Review, 27: 1-26. Duplaix N., 1980. Observations on the ecology and behavior of the giant river otter Pteron ura brasiliensis in Suriname. Revue Ecologique (Terre Vie), 34: 495- 620. Eisenberg J.F., 1989. Mammals of the Neotropics. Volume 1. The Northern Neotropics. Chicago: University of Chicago Press, Chicago, CHI. Hajek F. & Groenendijk J., 2006. Lobos del rio Madre de Dios. Lima: Ayuda para la vida silvestre amenazada. Sociedad Zoologica de Francfort Pern, Lima, Pern. Gandiwa E., 2012. Local knowledge and perceptions of animal population abundances by communities adjacent to the northern Gonarezhou National Park, Zimbabwe. Tropical Conservation Science, 5: 255- 269. Gonzalez J.A., 2003. Harvesting, local trade, and conser- vation of parrots in the Northeastern Peruvian Amazon. Biological Conservation, 114: 437-446. Groenendijk J., Hajek F., Isola S. & Schenk C., 2001. Giant otter project in Pern-field trip and activity re- port-2000. IUCN Otter Specialist Group Bulletin, 18: 20-27. IUCN, 2006. Red List of Threatened Species. Version 2015-4. . Downloaded on 28 January 2016. Kotchwar Logan M., Gerber B.D., Karpanty S.M., Justin S. & Rabenahy F.N., 2015. Assessing carnivore distri- bution from local knowledge across a human-domin- ated landscape in central-southeastern Madagascar. Animal Conservation, 18: 82-91. Myers N., 1988. Threatened biotas: “hotspots” in tropical forests. Environmentalist, 8: 187-208. Piana Renzo P., Del Aguila Chavez J. & Tang Tuesta M., 2003. Experiencias de manejo de paiche en cuatro comunidades de la Reserva Nacional Pacaya Samiria. In: Seminario Taller Internacional de Manejo de Paiche o Pirarucu, Iquitos, Peru. Alcantara F. & Montreuil V. (Eds.). Instituto de Investigaciones de la Amazonia Peruana (IIAP) and World Wildlife Foundation (WWF) - Russell E. Train Education for Nature Program, 29-44. Puertas P., Bodmer R., Parodi J.L., Aguila J. & Calle A., 2000. La importancia de la participacion comunitaria en los planes de manejo de fauna silvestre en el nor- oriente del Pern. Folia Amazonica, 11: 159-172. Recharte Uscamaita M. & Bodmer R., 20 10. Recovery of the endangered giant otter Pteronura brasiliensis on the Yavari-Mirin and Yavari Rivers: a success story for CITES. Oryx, 44: 83-88. Sanderson E.W., Jaiteh M., Levy M.A., RedfordK H., Wannebo A.V. & Woolmer G., 2002. The human foot- print and the last of the wild. Bioscience, 52: 891- 904. Scholte R, 2011. Towards understanding large mammal population declines in Africa's protected areas: A West-Central African perspective. Tropical Conserva- tion Science, 4: 1-11. Shostell J.M. & Ruiz-Garcia M., 2013. An introduction to neotropical carnivores. In: Molecular Population Genetics, Evolutionary Biology and Biological Conservation of the Neotropical Carnivores. Ruiz Garcia M. & Shostell J.M. (Eds.), Nova Science Publishers Inc., New York, NY, 1-45. Biodiversity Journal, 2016, 7 (1): 25-32 Patterns of Butterfly distribution in Alabama, USA (Lepidop- tera) Xiongwen Chen* &Tuo Feng Department of Biological and Environmental Sciences, Alabama A & M University, Normal, AL 35762, USA * Corresponding author, e-mail: xiongwen.chen@aamu.edu ABSTRACT Butterflies (Lepidoptera) are an iconic group of insects and are emphasized in ecological research and biodiversity conservation due to the role in ecological processes. Alabama (USA) has 139 species of butterflies in 6 families based on the previous field surveys. In this study the information from the previous field survey was analyzed with environmental information for the general patterns across 67 counties of Alabama. The results indicate that the counties with the higher butterfly species are mainly within the metropolitan areas; power-law relationship exists between average species number and occupied county number; there is higher number of butterfly species at counties with either the highest or the lowest forest coverage; there is positive correlation between latitude and butterfly species density; counties with the lowest or the highest species number usually have higher standard deviations in annual air temperature or precipitation; butterflies with a big distribution area do not have significantly bigger wing size in comparison to ones with a small distribution area; and with the increase of latitude, the average wing size of butterflies increases. The results provide new understanding for the butterfly distribution at a regional level. KEY WORDS Alabama; butterflies; climate; latitude; species number; wing size. Received 02.02.2015; accepted 21.03.2016; printed 30.03.2016 INTRODUCTION The Butterflies (Lepidoptera) play an important role in ecosystems and conduct ecological services (Tiple et al., 2006), such as pollination and herbi- vores. Butterflies are considered as good ecological indicators of the health of some terrestrial eco- systems (New, 1991; Thomas, 2005; Bonebrake et al., 2010). The beautiful color of butterflies and unique features also provide recreation resource to human society. Butterflies are greater sensitive than other taxonomic groups to reflect human disturb- ance (Thomas, 2005). Monitoring butterfly species at an area can indicate human mismanagement and pollution (Wilson, 1997). Due to climate change, altered land use (e.g., habitat loss), and pollutants (e.g., pesticides and herbicides), the butterflies are in declining, such as in Europe (van Swaay et al., 2006). The loss of native plants, which are food for leaf-eating caterpillars and nectar sipping adult butterflies, by the replacement of exotic invasive species has devastated butterflies. Butterflies are an iconic group of insects and are emphasized in eco- logy and biodiversity conservation. The state of Alabama (USA) has 139 species of butterflies in six families (Hesperiidae, Papilionidae, Pieridae, Lycaenidae, Riodinidae, and Nymphal- idae). The information of distribution, habitat, food, life history and wingspan for all 139 species is listed in the book “Butterflies of Alabama” based on the 26 Xiongwen Chen &Tuo Feng field records (Howell & Charny, 2010). This in- formation provides an opportunity for integrated study, such as analyzing patterns of butterfly distri- bution and uncovering the related factors. One of the important features of butterflies is their wingspan or body size. Body size is a key trait related to the life history of individuals, the wing size (a proxy for body size) of butterflies signi- ficantly decreased in response to warmer summers in high arctic area (Bowden et al., 2015). Based on the Bergmann’s rule, larger individuals occur at higher latitudes and in colder environments (Sand et al., 1995). Similarly, smaller adult size should be in higher temperatures or southern area. Although both Bergmann’s rule and the temperature-size rule predict larger individuals in colder environments, however, the opposite pattern also reported (Blanck- enhorn & Demont, 2004; Angilletta, 2009). Several ways were proposed that temperature may affect body size. Two mechanisms related to external temperat- ures may impact body size in different directions. First, the metabolic rates increase with warmer temperatures, organisms become smaller if they cannot offset energy losses under high metabolic costs. Second, rising temperatures in seasonal environ- ment make longer growing seasons, which may let organisms grow larger. The extended seasons could also low plant-food quality during late season (Awmack & Leather, 2002). Baguette & Stevens (2013) suggested that wingsize of butterflies is positively related to min- imum area requirements. Butterflies with big wing size should have a big distribution area. Host-range relationship may be primarily determined by eco- logical and population-genetic factors (Barrett & Heil, 2012). For example, generalists should be promoted by volatile host communities, while specialists should be favored in places where host communities are stable (Jaenike, 1990). This means that harsh and volatile climate in a temperate region could have more generalists and favorable and static climate have more specialists. For the distri- bution area, plants are food and habitats to butter- flies, forests harbor between 50% and 90% of Earth’s terrestrial species including diverse of plant species (World Resources Institute et al., 1992), there should have more butterfly species in forest areas than at less or none forest areas. It is also known that butterflies are sensitive to habitat fragmentation (Ockinger et al., 2010), so with the increased landscape fragmentation in one region, such as in a metropolitan area, butterfly species number may decrease. Therefore, the goal of this study is to use the collected butterfly inform- ation from Howell & Charny (2010) combined with climate and environmental information to indicate the general patterns of butterfly distribution in the state of Alabama and test the above hypotheses. The specific objectives include (i) distribution pattern of butterfly species along latitude; (ii) relationship between wing size of butterflies and latitude; (iii) relationship between butterfly species number and plant species number and forest cover at county level; and (iv) relationship between butterfly species number and urbanization at the county level. This study will provide understanding of the patterns of butterfly distribution in Alabama. MATERIAL AND METHODS Study area Alabama is located in the southern region of USA. and between the southern foothills of the Appalachian Mountain Range and the Gulf of Mexico. There are total 67 counties in Alabama (Fig. 1). Since the State of Alabama runs roughly from 31° to 35°N, the climate in the southern part is warmer than the northern part. Northern Alabama has a warm, humid, temperate climate, and the south has a subtropical climate. Summers are hot and humid with an average high temperature around 33°C; winters are typified by a series of cold fronts. The annual precipitation varies from 150 cm to 162 cm in the northern part and 180 cm to 195 cm in the southern part (Carter & Carter, 1984). Based the inventory data from Alabama Forestry Commission (www.forestry.state.al.us), 70% of the state is covered by forests. Due to mild climate and hetero- geneous landscape, Alabama has great species diversity. The county level is selected in this study because most data are only available at this level. Data Butterflies: the butterfly information is from the book of Howell & Charny (2010), which was based Patterns of Butterfly distribution in Alabama, USA (Lepidoptera) 27 on the year-round field observations from 2001 to 2009 by the authors, their students and colleagues. Photographic survey which is broadly applied for biodiversity research (e.g, McGrath, 2015) was conducted at each county. The spatial resolution of butterfly distribution is at county level, which means the distribution covers the entire county as long as this butterfly species is found at one loca- tion. More information can be found in Howell & Chamy (2010). In this study, the information of distribution and the average wing size is used. Climate: the climate information is from local weather stations in each county from 2001 to 2009. Plants and forest: the information of plant species diversity in each county of Alabama is from http://www.alabamaplants.com. The forest cover- age (%) in each county at that time is from Chen (2009). Human population: the human population at each county during the corresponding time period is obtained from Alabama Quick Facts at the USCensus Bureau (http://quickfacts.cencus.gov/ qfd/index). 1 11 -26 Number | 19 -49 I 50-79 0 30 60 120 Kilometers Li-i-i J . l.L-lJ Statistical method Standard deviation was used to characterize the fluctuation in air temperature and precipitation in each county. The commonly used least squares technique was used in correlation analysis and T- test of SAS (SAS Institute Inc., NC, USA.). The statistical test was considered significant at p<0.05. Data aggregation was applied when the statistical test on individual county data was not significant, but the trend might exist, such as the bin of [0, 10], [11, 20],... [70, 80] was applied for the rank of butterfly species number while testing power-law between the average species number and appeared county number. The butterfly species density in each county was estimated by the total butterfly species number /county area. RESULTS Jefferson County has 79 butterfly species, which is the highest number. The counties with the cat- egory of highest butterfly species (50-79) include Madison, Jackson, Tuscaloosa, Jefferson, Ribb, Shelby, and Baldwin (Fig. 1). These counties are Figure 1 . The butterfly distribution across the counties of Alabama (bold lines indicate metropolitan area). mainly within the metropolitan areas of Hunts- ville, Birmingham, and Mobile cities. There are six counties (Choctaw, Coffee, Crenshaw, Dale, Greene, and Lamar) without any butterflies or with veiy limited species number. There is a power-law re- lationship between the average of butterfly species number and appeared county number (Fig. 2). The relationship between county size and butterfly species number is not obvious (Fig. 3). The correlation between human population in each county and butterfly species is not significant (p> 0.05) (Fig. 4). The relationship between plant species number and butterfly species number among all the counties is not obvious (Fig. 5). There is higher number of butterfly species at areas with either the highest or the lowest forest coverage (Fig. 6). There is positive correlation between latitude and butterfly species density (Fig. 7). The correla- tion between the average annual air temperature or average annual precipitation and species density is not significant (p>0.05) (Fig. 8), but there is a general trend of decreased species density with 28 Xiongwen Chen &Tuo Feng Figure 2. The correlation between average butterfly species number and appeared county number. Figure 5. The relationship between plant species number and butterfly species number in counties. 90 80 ▼ 70 5 60 i so C S 40 20 10 fl *-♦— 4 ♦ 4 4 — $ — ♦♦ 4 4 444 4 4 V* ♦ 4 4 Tittit ♦ * 2 * ♦ Y ♦♦44 4 " — * 9.1 9.2 9.3 9.4 9.5 9.6 9.7 9.8 log (countYarea, m') Figure 3. The relationship between county size and butterfly species number. Figure 6. The relationship between forest coverage and butterfly species number in counties. Figure 4. The relationship between human population and butterfly species number among counties. Figure 7. The relationship between latitude and density of butterfly species. Patterns of Butterfly distribution in Alabama, USA (Lepidoptera) 29 Figure 8. The relationship between butterfly species number and average annual air temperature (a) and average annual precipitation (b). Figure 9. The relationship between butterfly species number and standard deviation of annual air temperature (a) and standard deviation of annual precipitation (b). Figure 10. The relationship between average wingsize of butterfly and the diameter of the distribution area. Figure 11. The relationship between latitude and butterfly wingsize. 30 Xiongwen Chen &Tuo Feng increased temperature or precipitation. There is a pattern that counties with the lowest or the highest species number have higher standard deviations in annual air temperature or precipitation (Fig. 9). The correlation between the average wingsize and diameter of distribution area at each county level is not significant (p>0.05). However, after the data aggregation in wingsize, there is a gen- eral trend between the average wingsize and the diameter of distribution area (Fig. 10). The aver- age wingsize of the broadly distributed species (or generalists) is 53. 7± 25.7 mm and 51.0 ± 23.7 mm for narrow distributed species (or specialists). The difference in wing size between generalists and specialists is not statistically significant (p> 0.05). With the increase of latitude, the average wingsize increases for all species polled over (Fig. 11). DISCUSSION AND CONCLUSIONS There are some patterns of butterfly distribution in Alabama after the integrated analysis with other information. Some counties have a high species number, but others have limited species. The power-law relationship between average species number and appeared county number is similar to those with plants and animals in California (Chen et al., 2006). The phenomena may be related to the spatial occupying process and tolerance of habitat for all the species, but the mechanism is not known. With the increase of county size in area, this does not necessary lead to the increase in the number of butterfly species, which means big counties may not have more butterfly species. The island biogeo- graphy theory does not apply to butterfly species here. The counties with higher number of butterfly species are mainly within these metropolitan areas (e.g., major cities of Birmingham, Huntsville and Mobile areas). It seems that the higher number of butterfly species is related to human population and land use change, although the correlation between butterfly species number and human population in each county is not significant. This is consistent to that (i) no obvious relationship between butterfly species number and plant species number among all counties; (ii) there is high species number at areas with either the lowest or highest forest coverage. After comparing the butterfly species diversity in urban, suburban and rural areas, Mukherjee et al. (2015) indicated that butterfly species diversity is related to landscape heterogeneity. Usually there is higher landscape heterogeneity at the metropolitan areas due to diverse vegetation pattern under different land uses from land owners, but relatively homogeneity landscape in urban and rural areas. Earlier studies suggested that butterfly diversity is attributed to plant species (Kuussaari et al., 2007). But in this study, there is no obvious correlation between plant species and butterfly species at county level. These butterfly species may only like some specific plants for hosting (Howell & Chamy, 2010 ). Usually in warmer area, such as tropical areas, there is higher species diversity. However, in this study the relationship between latitude and butterfly species is on the opposite. There is higher density of butterfly species in northern Alabama. This result is also consistent with that there is a general trend of decreased species density with increased tem- perature. The possible cause may be that the rule at continental (or global) level may not always work at a regional level. Some additional factors may attract to butterfly species diversity at a regional level. Also, in low latitude areas there are high species number as overall, but not necessary for butterfly species. The results in this study also identify that counties with large fluctuations in annual air temperature and precipitation have either the highest or the lowest species number of butterfly. Under the stable climate condition (e.g., lower standard deviation in annual temperature or pre- cipitation) there is an intermediate high number of butterfly species. The changing climate may provide more niche space for various butterfly species if they can tolerate. The degree to which phenotypic plasticity and adaptation ultimately play a role under this changing climate remains to be further studied (Bowden et al., 2015). Bergmann’s rule, describing the relation between latitudinal and body size, is confirmed in this study. Our res- ults indicate that the average wingsize of butter- fly increases with the increase of latitude in Alabama. There are generalists of butterfly with a large distribution area from the south to north and also several specialists with limited distribution in Ala- bama (such as only one county). But the sizes of their wingspans are not significantly different. This Patterns of Butterfly distribution in Alabama, USA (Lepidoptera) 31 result may indicate that butterfly species with big wingspans may not necessary show greater migra- tion capacity or the specialists may also be distrib- uted broadly if resource is suitable. The size of wingspan may not determine the fate of some specialist of butterflies under changing environment which was considered as venerable (Dapporto & Dennis, 2013). After analyzing the records of butterfly species and the environmental factors in Alabama, the emergent patterns at a regional level appear for the distribution of butterfly. The uneven distribution of butterfly species may be related to land use and climate fluctuations. The species diversity and body size related with latitude and temperature may provide helpful information for butterfly conserva- tion and mitigation under climate change. This study may provide a background map for study of butterfly distribution under environmental change (McGrath, 2015). Periodically monitoring the body size and distribution of butterfly species and other biodiversity may be necessary for sustainable regional development. ACKNOWLEDGMENTS This work was partially supported by the USD A National Institute of Food and Agriculture Mclntire Stennis project (1008643). REFERENCES Angilletta Jr M.J., 2009. Thermal adaptation: a theoreti- cal and empirical synthesis. Oxford University Press, Oxford. 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Carter E.A. & Carter V.G.S., 1984. Extreme weather history and climate atlas for Alabama. Strode Publishers, Huntsville, Alabama. Chen X., 2010. Trends of forest inventory data in Alabama, USA during the last seven decades. Forestry, 83: 517-526. Chen X., Li B.-L., Scott T. & Allen M.F., 2006. Tolerance analysis of habitat loss for multispecies conservation in western Riverside County, California, USA. Inter- national Journal of Biodiversity Science and Man- agement, 2: 87-96. Dapporto L. & Dennis R.L.H., 2013. The generalist- specialist continuum: Testing predictions for distri- bution and trends in British butterflies. Biological Conservation, 157: 229-236. Howell W.M. & Chamy V., 2010. Butterflies of Alabama. Pearson Learning Solutions, Boston, MA. Jaenike J., 1990. Host specialization in phytophagous insects. Annual Review of Ecology and Systematics, 21: 243-273. Kuussaari M., Heliola J., Luoto M. & Poyry J., 2007. Determinants of local species richness of diurnal Lepidoptera in boreal agricultural landscapes. Agri- culture, Ecosystems & Environment, 122: 366-376. Mukherjee S., Banerjee S., Saha G.K., Basu P. & Aditya G., 2015. Butterfly diversity in Kolkata, India: An appraisal for conservation Management. Journal of Asia-Pacific Biodiversity, 8: 210-221. McGrath P.F. 201 5. A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy. Biodiversity Journal, 6: 53-72. New T.R., 1991. Butterfly conservation. Oxford Univer- sity Press, Melbourne. Ockinger E., Schweiger O., Crist T.O., Debinski D.M., Krauss J., Kuussaari M., Petersen J.D., Poyry J., Set- tele J., Summerville K.S. & Bommarco R., 2010. Life history traits predict species responses to habitat area and isolation: a crosscontinental synthesis. Ecology Letters, 13: 969-979. Sand H.K., Cederlund G.R. & Danell K., 1995. Geo- graphical and latitudinal variation in growth patterns and adult body size of Swedish moose (Alcesalces). Oecologia, 102: 433-442. Thomas J.A., 2005. Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups. Philosophical Transactions of the Royal Society B, 360: 339-357. Tiple A.D., Deshmukh V.P. & Dennis R.L.H., 2006. Factors influencing nectar plant resource visits by 32 Xiongwen Chen &Tuo Feng butterflies on a university campus: implications for conservation. Nota Lepidopteralogica, 28: 213-224. Van Swaay C., Warren M. & Lot's G., 2006. Biotope use and trends of European butterflies. Journal of Insect Conservation, 10: 189-209. Wilson E.O., 1997. Introduction. In: Reaka-Kudla M.L., Wilson D.E. & Wilson E.O. (Eds.), Biodiversity II. Henry Press, Washington, D.C., pp. 1-3. World Resources Institute, United Nations Environment Program, United Nations Development Program, 1992. World Resources 1992-93: a Report. Oxford University Press, New York. Biodiversity Journal, 2016, 7 (1): 33-38 First record of a Humpback Whale Megaptera novaeangliae (Borowski, 1 78 1) in theTyrrhenian Sea (Cetacea Balaenopte- ridae) Nicola Maio 1 *, Vincenzo Maione 2 & Riccardo Sgammato 2 'Dipartimento di Biologia, Complesso Universitario di Monte Sant’Angelo, Universita degli Studi di Napoli Federico II, Edificio 7, via Cinthia 26, 80126 Napoli, Italy. 2 Centro Sub Campi Flegrei, Via Miliscola 165, 80078, Pozzuoli, Frazione Lucrino, Napoli, Italy. ^Corresponding author: e-mail: nicomaio@unina.it ABSTRACT It is reported the sighting of a Humpback Whale Megaptera novaeangliae (Borowski, 1781) (Cetacea Balaenopteridae) in the Gulf of Pozzuoli, near the coast of Baia (Bacoli, Napoli, Campania, Southern Italy). This record represents the first in the Tyrrenian Sea, the eighth in the Italian Seas and the twenty-fourth in the Mediterranean Sea. KEY WORDS Megaptera novaeangliae', Humpback Whale; sighting; Tyrrhenian Sea. Received 19.02.2016; accepted 07.03.2016; printed 30.03.2016 INTRODUCTION The Humpback Whale, Megaptera novaeangliae (Borowski, 1781) (Order Cetacea, Suborder Mys- ticeti, Family Balaenopteridae) is a cosmopolitan species widely distributed and far-ranging migrant, found in both hemispheres and in all the major ocean basins. During the winter, at the period of mating and calving grounds, all the populations mi- grate to tropical waters, usually near continental coastlines or island groups; during spring, summer and autumn they move to productive colder waters in temperate and high latitudes, where most of the feeding takes place. In the North Atlantic, during the summer the Humpback Whale ranges from the Gulf of Maine in the West and Ireland in the East, and in the North but not into the pack ice; the northern extent of the Humpback's range includes the Barents Sea, Greenland Sea and Davis Strait (but not the Canadian Arctic), where they occur mainly in specific feeding areas. During the winter, the majority of whales migrate to wintering grounds in the West Indies, and an apparently small number use breeding areas around the Cape Verde Islands. In the Mediterranean Sea, the Humpback Whale is not regularly present; in fact it is considered as an irregular or occasional “visitor species”, accord- ing to the Reports of Agreement on the Conserva- tion of Cetaceans of the Black Sea, Mediterranean Sea and Contiguous Atlantic Area (ACCOBAMS), entering the region from the Strait of Gibraltar (Reeves & Notarbartolo di Sciara, 2006; Notarbar- tolo di Sciara & Birkun, 2010; Cagnolaro et al., 2015). Since 1990 the number of observations has increased and the range of sighting locations has expanded so as to include both basins of the Mediter- ranean Sea (Frantzis et al., 2004). Humpback Whale, Megaptera novaeangliae , is well known for his long pectoral fms, which can be up to 4.6 meters in length. The dorsal fin is 34 Nicola Maio et alii variable in size and shape, from small triangular knob to larger sickle-shaped, placed nearly two- thirds along back. Head and body are black or grey, white on throat and belly. The adult can measure up to 17 m. MATERIAL AND METHODS We take into consideration the sighting of one individual photographed in the Bay of Pozzuoli; the sighting occurred from the Aragonese Castle of Baia (District of Bacoli Municipality, Province of Naples) at about 70 m of height. The camera equip- ment consisted of a Digital single-lens reflex camera Canon EOS 650D with 75-300mmEF-S lens mounted. RESULTS AND DISCUSSION Here we report the sighting of a Humpback Figure 1. The locations of sightings of Humpback Whales, Megaptera novaeangliae, in the Italian Seas. First record of a HumpbackWhale Megaptera novaeangliae in the Tyrrhenian Sea (Cetacea Balaenopteridae) 35 Figures 2-4. Humpback Whale, Megaptera novaeangliae, recorded near Baia, in the Bay of Pozzuoli, apparently in good conditions (Photos by R. Sgammato). 36 Nicola Maio et alii DATE LOCATION EVENT ANIMALS, SIZE SOURCE AND NOTES 1998, 24 January Gulf of Oristano (Sardinia) Sardinian Sea Sighting 1 (7-9 m) (Lrantzis et al., 2004) 2002, 4 August Senigallia (Province of Ancona, Marche) Adriatic Sea Sighting 1 (Affronte et al., 2003) 2004, 2 April Syracuse (Sicily) Ionian Sea Accidentally by-caught and released 1 (about 10 m) Centro Studi Cetacei, 2006 2010, 26-28 August Eastern Ligurian Sea: Versilia (Prov. of Lucca, Tuscany) Sestri Levante (Prov. of Genoa, Liguria) Repeated sightings of one individual 1 (about 10- 13m) (Cagnolaro et al., 2015) 2011,24 March Near Savona (Liguria) Ligurian Sea Sighting (Cagnolaro et al., 2015) 2013, 12 March Lampedusa Island (Sicily) Sicily Channel Sighting of one indivi- dual already observed in Trench Ligurian Sea 1 (8-9 m) (Panigada et al., 2014) 2013, August Ligurian Sea Sighting of the same individual of Lampedusa 1 (8-9 m) (Panigada et al., 2014) 2015, 10 December Baia, Bay of Pozzuoli (Province of Naples, Campania) Tyrrenian Sea Sighting 1 Present work Table 1. Reports concerning specimens of Humpback Whale, Megaptera novaeangliae, recorded in the Italian seas. Whale, Megaptera novaeangliae, in the Bay of Poz- zuoli near Baia, a District of Bacoli Municipality (Province of Naples, Campania Region) occurred on 10 December 2015. The animal has been obser- ved near the coast at a depth of about six meters, it was approximately 8-9 meters long (probably a ju- venile) and with the uppersides of both pectoral fins of white color, apparently in good conditions (Figs. 2-4). This is the first documented record of a Hum- pback Whale in the Tyrrenian Sea, and the first si- ghting for Campania Region (Maio & Quercia, 2006; Maio et al., 2012). Our finding suggests that the Tyrrenian waters offer suitable habitats also for this species. Since 1885, 24 records (16 sightings of which four with two individuals, three strandings and 5 by-caught individuals) have been reported from dif- ferent locations across the Mediterranean basin. All individuals, ranging between 7 and 12 meters, were estimated to be 2-3 years old juveniles (Panigada et al., 2014). The first occurrence in the Mediterranean Sea was a juvenile caught in 1885 off 15 km West of Toulon (France) (Pouchet, 1885; Beauregard, 1885; VanBeneden, 1889; Aguilar, 1989). Occurrences of Flumpback Whales, Megaptera novaeangliae, are extremely rare in the Italian Seas being known only six sightings and one captures of single specimens. Date, location and size are given in Table 1 . The first occurrence was of a 7-9 m long individual reported in the Gulf of Oristano (Sardinia), in January 1998 (Frantzis et al., 2004). The last sighting was an individual approxim- ately 8-9 meters long, observed in three different First record of a HumpbackWhale Megaptera novaeangliae in the Tyrrhenian Sea (Cetacea Balaenopteridae) 37 locations: the first time it was observed in the French Liguria Sea, NW Mediterranean, in June 2012; then, the same animal was re-sighted off Lampedusa Island, Sicily Channel, in March 2013 over 1,000 km away in a straight line from the previous location and again in August 2013, in the “Italian” Ligurian Sea (Panigada et al., 2014). No specimens from Mediterranean Sea are preserved in Italian museums (Cagnolaro et al., 2014). The Humpback Whale, Megaptera novaeangliae , is a species listed in the Appendix I of CITES, and it is considered an “Endangered or threatened species” in the Annex II of the Barcelona Conven- tion for Protection against Pollution in the Mediter- ranean Sea. It is also included in the Appendix II of the Bern Convention on the Conservation of European Wildlife and Natural Habitats, con- sidered as “Strictly protected fauna species”, and is a “species in need of strict protection” in European Union by the Annex IV of the Council Directive 92/43/EEC of May 21thl992 on the conservation of natural habitats and of wild fauna and flora, known as “Habitats Directive”. Further- more the species is classified as “Least Concern” on the IUCN Red List of Threatned Species (vers. 2015.4) (Reilly et al., 2008). AKNOWLEDGEMENT We wish to thank Gennaro Bianco (University of Naples “Parthenope”, Italy), Lucia Borrelli, Elena Confalone and Roberta De Stasio (Univer- sity of Naples Federico II, Italy), Gianfranco Pol- laro (Centro Studi Ecosistemi Mediterranei, Pollica, Salerno, Italy). REFERENCES Affronte M., Stanzani L.A. & Stanzani G., 2003. First re- cord of the humpback whale, Megaptera novaeangliae (Borowski, 1781), from the Adriatic Sea. Annales, Series Historia Naturalis, 13: 51-54. Aguilar A., 1989. A record of two Humpback Whales, Megaptera novaeangliae, in the Western Mediter- ranean Sea. Marine Mammal Science, 5: 306-309. doi: 10. 1 1 1 1/j. 1 748-7692. 1989. tb00344.x Beauregard H., 1885. Note sur une Megaptere echouee au Bmsc pres Toulon. Seances du 19 Decembre 1885. Comptes Rendus Hebdomadaires des Seances et Memories de la Societe de Biologie, Paris [Compt. Rend. Mem. Soc. Biol. Paris], 37 (tome 2, serie 8): 753. Cagnolaro L., Cozzi B., Notarbartolo di Sciara G. & Podesta M. (Eds.), 2015. Fauna d’ltalia Vol. XLIX. Mammalia IV. Cetacea. Calderini, Bologna, 390 pp. Cagnolaro L., Maio N. & Vomero V. (Eds.), 2014. The Cetacean collections of italian museums. First part (living Cetaceans). Museologia Scientifica. Memorie, 12: 1-420. Centro Studi Cetacei Onlus & Museo Civico di Storia Naturale di Milano, 2006. Cetacei spiaggiati lungo le coste italiane. XIX. Rendiconto 2004. Atti della Sociea italiana di Scienze naturali Museo civico di Storia naturale di Milano, 147: 147-157. Frantzis A., Nikolaou O., Bompar J.-M. & Cammedda A., 2004. Humpbackwhale (Megaptera novaeangliae ) occurrence in the Mediterranean Sea. The Journal of Cetacean Research Management, 6: 25-28. Maio N., Pollaro F., Di Nocera F., De Carlo E. & Galiero G., 2012. Cetacei spiaggiati lungo le coste della Campania dal 2006 al 2011 (Mammalia: Cetacea). Atti della Sociea italiana di Scienze naturali Museo civico di Storia naturale di Milano, 153: 241-255. Maio N. & Quercia F., 2006. Cetacei spiaggiati lungo il litorale campano: ricerca e conservazione. In: Gugliemi R. & Nappi A. (Eds.). Atti Convegno: “Fa Natura in Campania: aspetti biotici e abiotici”. Napoli, 18 novembre 2004, 158-164. Notarbartolo di Sciara G. & Birkun A. Jr., 2010. Con- serving whales, dolphins and porpoises in the Mediter- ranean and Black Seas: an ACCOBAMS status re- port, 2010. ACCOBAMS, Monaco. 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The IUCN Red Fist of Threatened Species 2008: e.T13006A3405371. http://dx.doi.org/10.2305/IUCN.UK.2008.RFTS.T13 006A3405371.en. [last visit date: 18.02.2016]. 38 Nicola Maio et alii Van Beneden P.J., 1889. Histoire naturelle des Cetaces des Sciences, des Lettre set des Beaux- Arts de des mers d’Europe. Imprimeur de Academie Royale Belgique, Bruxelles, 1-6. Biodiversity Journal, 2016, 7 (1): 39-50 Systematic account of Orthoptera fauna of Bastar district, Chhattisgarh, India Sunil Kumar Gupta Zoological Survey of India, Prani Vigyan Bhawan, 'M' Block, New Alipore, Kolkata, 700053 West Bengal, India; e-mail: skumarento@gmail.com ABSTRACT A faunistic survey in Bastar district, Chhattisgarh (India) revealed 52 species belonging to 45 genera, 8 families, including five species which are new record to the Orthoptera fauna of Chhattisgarh: Calliptamus barbarus bar bar us (Costa, 1836), Ceracris fasciata (Brunner von Wattenwyl, 1893), Oedaleus senegalensis (Krauss, 1877), A u larches miliaris miliaris (Linnaeus, 1758), and Loxoblemmus haani Saussure, 1877. KEY WORDS Distribution; Bastar; Orthoptera; Chhattisgarh. Received 23.02.2016; accepted 08.03.2016; printed 30.03.2016 INTRODUCTION The major works on Orthopteran fauna of India were published by Kirby (1914) and Chopard (1969). Notable papers on Orthoptera fauna of Chhattisgarh state were also done by Dwivedi (1978, 1990), Dixit & Sinha (1982), Agrawal & Sinha (1987), Chandra & Gupta (2005), Chandra et al. (2007), Gupta et al. (2008), Gupta & Chandra (2010) and Gupta & Shishodia (2014), but so far no comprehensive account on Orthoptera of Bastar is available a part from a few exceptions including: Chopard (1970) who described Arachnomimus sub- alatus Chopard, 1970 and Sinha & Agrawal (1973) who described Kempiola shankari (Sinha et Agrawal, 1973) both from the same locality, i.e. Kutums war cave. Shishodia, (1995) reported 15 species belong- ing 15 genera under 6 families from Indravati Tiger Reserve, Bastar. Shishodia (2000) reported 77 species of crickets and grasshoppers from Bastar. MATERIAL AND METHODS A total of 514 specimens representing 52 species belonging 45 genera under 8 families viz. Ac- rididae 28 species 21 genera, Pyrgomorphidae 4 species 4 genera, Tetrigidae 4 species 4 genera, Tridactylidae 1 species and 1 genus, Gryllidae 6 species 6 genera, Trigonidiidae 2 species 2 genera, Gryllotalpidae 1 species 1 genus, Tettigoniidae 6 species 6 genera, are known from Bastar district of Chhattisgarh. Of these, five species are reported for the first time. In Table 1 are showen coordinates of collection localities. The species recorded for the first time are marked with an asterisk (*). All specimens are preserved in collection R.P. Gupta & co-workers collection. SYSTEMATIC Order ORTHOPTERA Suborder CAELIFERA Superfamily ACRIDOIDEA Family ACRIDIDAE Subfamily ACRID IN AE Genus Acrida Linnaeus, 1758 40 Sunil Kumar Gupta S. No. Site Latitude N Longitude E Altitude m 1 Asna Village 19°7T5.4" 82°01' 20.9" 539 2 Amaguda 19°9'45.4" 82°0T5.1" 553 3 Bhanpuri 19°19T7.4" 81°51T7.0" 514 4 Bhatiguda Village 19°2'53.4" 82°3'3.5" 515 5 Belguda Village 19°13'03" 81°58'55.1" 552 6 Chidaipadar 19°10T.7" 81°58T9.9" 543 7 Dongaghatpara 19°00'28.5" 81°05'08" 485 8 Erikpal Village 19°07T7.9" 82°03'34.9" 542 9 Gariya bahar river 19°4'53.2" 82°3T.9" 547 10 Hathguda 19°5'45.6" 82°3' 9.7" 561 11 Jagdalpur City 19°4'33.4" 82°1’51.7" 478 12 Jeeragaon 19°2'7.9" 82°9'39.1" 563 13 Kalcha 19°6'36.8" 82°6T8.9" 559 14 Kohkapal 19°8'32.1" 82°6'21.4" 562 15 Kolchur 19°10'5.8" 81°57'31.9" 555 16 Kopaguda Village 19°3'34.7" 82°6'43.1" 600 17 Kotamsar 18°52'45" 81°55'21.1" 487 18 Kurandi 19°1'49.5" 82°6T3.1" 578 19 Machkote range 19°0'52.4" 82°8'2.3" 555 20 Malgaon 19°8'6.9" 82°4'47.9" 551 21 Magedha 19°46'0.4" 81°53T6.9" 592 22 Makdi FRH 19°46'22.3" 81°54T2.8" 671 23 Mongrapal Village 19°H'26.9" 81°59'27.1" 572 24 Nakaguda Village 19°10'7.8" 81°2'47.4" 579 25 Neganar Village 19°12T.7" 81°1'3.3" 488 26 Piplavand 19°19'24.5" 81°55'39.2" 513 27 Pushpal 18°15'23.5" 82°4'53.2" 584 28 Rampal 19°13'39.9" 82°00'41.5" 599 29 Sonarpal beat 19°18'37.5" 81°51'51.5" 486 30 Taraguda 1909 - 25 " 82°6T7.1" 554 31 Tiwasguda 19°10'3" 82°2'48.5" 579 32 Ulnar 19°10'20.3" 82°7'28" 568 33 Umargaon Village 19°10'40.2" 82°1'36.2" 568 Table 1. Coordinates of collection localities of Bastar district, Chhattisgarh (India). Acrida exaltata (Walker, 1859) Truxalis exaltata - Walker, 1859: 222 Acrida exaltata - Dey & Hazra, 2003: 24 Examined material. Chhattisgarh; Bastar, Malgaon, 23.XI.2011, 1 male; 18.IV.2012, 1 male; Belaguda, 16.1.2012, 1 male; Jhariya Bahara Nala, 20. 111. 2012, 3 males; Kurandi, 23.III.2012, 2 males and 2 females; Erikpal Village, 24.11.2012, 1 male. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. 2. Acrida gigantea (Herbst, 1786) Truxalis gigantea - Herbst, 1786: 191 Acrida gigantea - Joshi et al., 2004: 71 Examined material. Chhattisgarh; Bastar, Rampal Village, 19.1.2012, 1 female; Bhanpuri, 20.X.2011, 2 females; Neganar Village, 4.1.2012, 1 male; Dongraghat para, 6.II.2012, 1 male; Jag- dalpur city, 13.11.2012, 1 male; Taraguda, 13.11.2012, 4 females; Ericpal Village, 25.11.2012, 1 female; Malgaon, 9.III.2012, 1 female; 10. 111. 2012, 1 female; Kohkapal, 14.III.2012, 2 females; Kumndi, 23.III.2012, 1 male and 1 female; Gariya Bahar river, 24.III.2012, 1 female; Machkote Range, 7.VI.2012, 2 males and 2 fe- males; Kopaguda, 22.V.2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. 3. Acrida turrita (Linnaeus, 1758) Gryllus ( Acrida ) turritus - Linnaeus, 1758: 427 Gryllus {Acrida) nasutus - Linnaeus, 1764: 118 Acrida turrita - Kirby, 1914: 98 Examined material. Chhattisgarh; Bastar, Mangra para, 7.1.2012, 1 male and 1 female; Be- laguda, 18.1.2012, 1 male; Dongaghat para, 7.11.2012, 1 female; 8.II.2012, 1 female; Malgaon, 7. II. 2012, 2 males; 10.III.2012, 1 male and 2 fe- males; Erikpal Village, 24.11.2012, 1 female; Kurandi, 23.III.2012, 1 male; Kolchur, 18.IV.2012, 1 male; 7.VI.2012, 1 male; Kopaguda, 22.V.2012, 1 female. Distribution in Chhattisgarh. Bastar and Raipur. Genus Phlaeoba Stal, 1860 Systematic account of Orthoptera fauna of Bastar district, Chhattisgarh, India 41 4. Phlaeoba infumata Brunner, 1893 Phlaeoba infumata - Brunner, 1893: 124 Phlaeoba infumata - Dey & Hazra, 2003: 25. Examined material. Chhattisgarh; Bastar, Sonarpal Beat, 17.X.2011, 1 male; Neganar Village, 4.1.2012, 1 male; Nakaguda Village, 24.1.2012, 1 female; Malgaon, 9.II.2012, 1 female; 10.III.2012 1 male; Erickpal Village, 24.11.2012, 1 male; Kohkapal, 14.III.2012, 3 males and 4 females; Kalcha, 24.IV.2012, 2 males; 18.VI.2012, 1 male; Machkote range, 7.VI.2012, 1 male and 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. 5. Phlaeoba panteli Bolivar, 1902 Phlaeoba panteli - Bolivar, 1902: 589 Phleoba panteli - Dey & Hazra, 2003: 27 Examined material. Chhattisgarh; Bastar, Jagdalpur range, 29.VIII.2011, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily CALLIPTAMINAE Genus Calliptamus Audinet-Serville, 1831 6. Calliptamus barbarus barbarus (Costa, 1 836) (*) Acridium barbarum - Costa, 1836: 13 Caloptenopsis punctata - Kirby, 1914: 208 Calliptamus barbarus barbarus - Massa, 2009: 81 Examined material. Chhattisarh; Bastar, Amaguda, 24.VIII.2011, 1 female. Distribution in Chhattisgarh. Bastar. Remark. New record from Chhattisgarh state. Subfamily CATANTOPINAE Genus Choroedocus Bolivar 1914 7. Choroedocus illustris (Walker, 1870) Heteracris illustris - Walker, 1870: 662, 663 Chroedocus illustris - Uvarov, 1921a: 109 Examined material. Chhattisgarh; Bastar, Jagdalpur range, 19.VIII.2011, 1 female. Distribution in Chhattisgarh. Bastar. 8 . Diabolocatantops innotabilis (Walker, 1870) Acridium innotabile - Walker, 1870: 629 Diabolocatantops innotabilis - Jago, 1984: 371 Examined material. Chhattisgarh; Bastar, Jag- dalpur range, 29.VIII.2011, 2 females; Neganar Vil- lage, 4.1.2012, 1 female; Mograpal Village, 6.1.2012, 2 females; Chidaipadar, 20.1.2012, 1 fe- male; Asna Village, 2.II.2012, 1 female; 4.II.2012, 2 females; Erikpal Village, 24.11.2012, 1 male; Malgaon, 9.III.2012, 2 males and 1 female; Kohkapal, 14.III.2012, 2 females; Gariya bahar river, 22.III.2012, 1 female; Machkote range, 7. VI. 2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Packyacris Uvarov, 1923 9. Pachyacris vinosa (Walker, 1870) Acridium vinosum - Walker, 1870: 587 Pachyacris vinosa - Shishodia & Dey, 2006: 107 Examined material. Chhattisgrah; Bastar, Makdi range, 8.XI.2011, 1 male. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Stenocatantops Dirsh et Uvarov, 1953 10. Stenocatantops splendens (Thunberg, 1815) Gryllus splendens - Thunberg, 1815: 236 Stenocatantops splendens - Shishodia, 2000: 63 Examined material. Chhattisgrah; Bastar, Mograpal Village, 7.1.2012, 1 males and 1 female; Asna Village, 4.II.2012, 1 female; Malgaon, 9. 111. 2012, 1 male; 10.III.2012, lmale and 3 fe- males; Kohkapal, 14.III.2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Xenocatantops Dirsh et Uvarov, 1953 11. Xenocatantops humilis humilis (Audinet- Serville, 1839) Acridium humile - Audinet-Serville, 1839: 662 Xenocatantops humilis humilis - Shishodia, 2000: 62 42 Sunil Kumar Gupta Examined material. Chhattisgarh; Bastar, MakdiPond, 9.IX.2011, 1 female; 7.VI.2012, 1 fe- male; Mograpal Village, 7.1.2012, 1 male; Malgaon, 9. III. 2012, 1 male; 9.XII.2012, 1 female; Ko- hkapal, 14.III.2012, 1 male; Kurundi, 23.III.2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. 12 . Xenocatantops karnyi (Kirby, 1910) Catantops karnyi - Kirby, 1910: 483 Xenocatantops karnyi - Shishodia, 2000: 62 Examined material. Chhattisgarh; Bastar, Belguda Village, 16.1.2012, 1 female; 18.1.2012, 1 female; Dongaghatpara, 7.II.2012, 1 female; Amaguda, 2.III.2012, 1 male; Malgaon, 10.III.2012, 1 male and 1 female; Kohlcapal, 14.III.2012, 1 male and 1 female; Jeeragaon, 26.III.2012, 1 female. Distribution in Chhattisgarh. Bastar and Raipur. Subfamily COPTACRID1NAE Genus Eucoptacra Bolivar, 1902 13. Eucoptacra praemorsa (Walker, 1870) Acridium saturatum - Walker, 1870: 628 Eucoptacra saturata - Uvarov, 1921b: 503 Eucoptacra praemorsa - Tandon, 1976: 10 Examined material. Chhattisgarh; Bastar, Mal- gaon, 10.III.2012, 1 female; Taraguda, 16.IV.20 12, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily CYRTACANTHACRIDINAE Genus Cyrtacanthacris Walker, 1870 14. Cyrtacanthacris tatarica (Linnaeus, 1758) Gryllus locusta tataricus - Linnaeus, 1758: 432 Cyrtacanthacris tatarica - Shishodia, 2000: 58 Examined material. Chhattisgarh; Bastar, Amaguda, 23.VIII.2011, 1 female; 24.VIII.2011, 1 male. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily EYEPREPOCNEMIDINAE Genus Tylotropidius Stal, 1860 15. Tylotropidius varicornis (Walker, 1870) Heteracris varicornis - Walker, 1870: 667 Tylotropidius varicornis - Shishodia, 2000: 60 Examined material. Chhattisgarh; Bastar, Belaguda, 18.1.2012, 1 male; Jagdalpur range, 15. VII. 2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily GOMPHOCERINAE Genus Leva Bolivar, 1909 16. Leva indica (Bolivar, 1902) Gymnobothrus indicus - Bolivar, 1902: 596 Leva cruciata - Bolivar, 1914: 65 Leva indica - Jago, 1996: 94 Examined material. Chhattisgarh; Bastar, Jag- dalpur range, 24.VIII.2011, 1 male; Malgaon, 23.XI.2011, 2 males and 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily HEMIACRIDINAE Genus C/onacris Uvarov, 1943 17. Clonacris kirbyi (Finot, 1903) Euthymia kirbyi - Finot, 1903: 622-629 Clonacris kirbyi - Tandon, 1976: 3 Examined material. Chhattisgarh; Bastar, Nakaguda, 19.1.2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily OEDIPODINAE Genus A iolopus Fieber, 1853 18 . Aiolopus thalassinus tamulus (Fabricius, 1798) Gryllus tamulus - Fabricius, 1798: 195 Aiolopus thalassinus tamulus - Shishodia, 2000: 49 Examined material. Chhattisgarh; Bastar, Mal- gaon, 23.XI.2011, 2 males and 1 female; 9.III.2012, 1 female; 1 0.III.20 12, 1 male and 1 female; Jag- Systematic account of Orthoptera fauna of Bastar district, Chhattisgarh, India 43 dalpur city, 13.11.2012, 1 female; Ericpal Village, 24.11.2012, 1 female; Kohkapal, 14.III.2012, 3 males and 2 females; Machkote Range, 7.VI.2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Ceracris Walker, 1870 19. Ceracris fasciata (Brunner von Wattenwyl, 1893) (*) Parapleurus fasciata - Brunner von Wattenwyl, 1893: 127 Rammeacris gracilis - Willemse, 1951: 66 Ceracris fasciata - Ingrisch, 1989: 235 Examined material. Chhattisgarh; Bastar, Jag- dalpur range, 29.VIII.2011, 2 females. Distribution in Chhattisgarh. Bastar. Remark. New record from Chhattisgarh state. 20. Ceracris nigricornis nigricornis Walker, 1870 Ceracris nigricornis - Walker, 1870: 791 Ceracris nigricornis - Kirby, 1914: 110 Examined material. Chhattisgarh; Bastar, Taraguda, 16.IV.20 12, 1 male. Distribution in Chhattisgarh. Bastar, Bilaspur, Kabirdham, Raipur. Genus Gastrimargus Saussure, 1884 21. Gastrimargus africanus africanus (Saus- sure, 1888) Oedaleus ( Gastrimargus ) marmoratus var. Afric- anus - Saussure, 1888: 39 Gastrimargus africanus africanus - Shishodia, 2000: 51 Examined material. Chhattisgarh; Bastar, Ranker, 27.VII.2011, 2 females; Amaguda, 25.VIII.2011, 1 female; Jagdalpur, 29. VIII.201 1,2 fe- males; Asna Village, 2.II.2012, 1 female; Machkote range, 7.VI.2012, 1 male and 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Kabirdham, Raipur. Genus Morphacris Walker, 1870 22 . Morphacris fasciata (Thunberg, 1815) Gryllus fasciatus - Thunberg, 1815: 230 Morphacris fasciata sulcata - Shishodia, 2000: 50 Examined material. Chhattisgarh; Bastar, Jagdalpur city, 29.III.2012, 1 male; Hathguda, 29. 111. 2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Oedaleus Fieber, 1853 23. Oedaleus abruptus (Thunberg, 1815) Gryllus abruptus - Thunberg, 1815: 233 Oedaleus abruptus - Ritchie, 1981: 104 Examined material. Chhattisgarh; Bastar, Chitrakot, 25.VIII.2011, 1 female; Nandpur Beat, 20.X.2011, 1 female; Pipalvond, 22.X.2011, 1 fe- male; Ulnar, 22.XI.20 11,1 female; Malegaon, 23. XI.2011, 3 females; 9.III.2012, 1 male and 2 fe- males; 1 0.III.20 12, 8 males and 1 female; Neganar Village, 5.1.2012, 1 female; Mangrapara, 6.1.2012, 1 male; Mograpal Village, 7.1.2012, 1 male and 2 females; Belguda Village, 16.1.2012, 1 female; Rampal, 19.1.2012, 1 female; Asna Village, 2.11.2012, 1 female; 4.II.2012, 2 males; Dongaghat- para, 7. II. 2012, 2 females; Erikpal Village, 24.11.2012, 1 female); 25.11.2012, 3 females; Gariya bahar river, 24.III.2012, 1 female; Taraguda, 16.IV.2012, 1 female; Kopaguda, 22.V.2012, 1 male; Machkote range, 7.VI.2012, 1 male; Hathguda, 29.XII.20 12, 2 males. Distribution in Chhattisgarh. Bastar, Kabird- ham Raipur. 24. Oedaleus senegalensis (Krauss, 1877) (*) Pachytylus senegalensis - Krauss, 1877: 56 Oedaleus senegalensis - Ritchie, 1981: 94 Examined material. Chhattisgarh; Bastar, Kohkapal, 14.III.2012, 1 female. Distribution in Chhattisgarh. Bastar. Remark. New record from Chhattisgarh State. Genus Trilophidia Stal, 1873 25. Trilophidia annulata (Thunberg, 1815) 44 Sunil Kumar Gupta Gryllus annulatus - Thunberg, 1815: 234 Trilophidia annulata - Shishodia, 2000: 52 Examined material. Chhattisgarh; Bastar, Ulnar, 22.XI.2011, 1 male; Malgaon, 23.XI.2011, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Kabirdham, Raipur. Subfamily OXYINAE Genus Oxya Audinet-Serville, 1831 26. Oxya hyla hyla Audinet-Serville, 1831 Oxya hyla - Audinet-Serville, 1831: 287 Oxya hyla hyla - Shishodia, 2000: 55 Examined material. Chhattisgarh; Bastar, Sonarpal Beat, 17.X.2011, 1 male and 1 female; Ericpal Village, 24.11.2012, 2 females; Mal- gaon, 9.III.2012, 5 females; Kohkapal Village, 14. 111. 2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Kabirdham, Raipur. Subfamily SPATHOSTERNINAE Genus Spathosternum Krauss, 1877 27. Spathosternum prasiniferum prasiniferum (Walker, 1871) Heteracris prasinifera - Walker, 1871: 65 Spathosternum prasiniferum prasiniferum - Shisho- dia, 2000: 53 Examined material. Chhattisgarh; Bastar, Chitrkote, 23.VIII.2011, 1 male; Nandpur Beat, 20. X.2011, 1 female; Sonarpal Beat, 17.X.2011, 3 males and 1 female; Bhanpuri, 19.X.2011, 1 male; 21. X.2011, 1 female; Asna, 1.XI.2011, 2 females; 4.H.2012, 2 females; Makdi Pond, 9.XI.2011, 2 males and 3 females; Mageda, 9.XI.2011, 1 female; Makdi Range, 11.XI.2011, 1 female; Malgaon, 23.XI.2011, 8 females; 7.III.2012, 1 male; 9.III.2012, 1 male and 4 females; 10.III.2012, 2 males and 5 females; Mograpal Village, 6.1.2012, 1 female; 7.1.2012, 1 fe- male; Belguda Village, 16.1.2012, 1 female; 18.1.2012, 1 female; Rampal Village, 19.1.2012, 1 male; Tiwasguda, 23.1.2012, 1 male; Dongraghat Para, 7.II.2012, 1 female; Taraguda, 13.11.2012, 2 fe- males; 12.III.20 12, 2 females; Kohkapal, 14.11.2012, 1 male; Erikpal Village, 24.11.2012, 1 male; 25.11.2012. 1 male; 10.III.2012, 1 male and 4 females; Kohkapal, 14.III.2012, 1 male and 1 female; Ulner Village, 16.III.2012, 1 male and 1 female; Gariya bahar river, 20.III.2012, 2 males and 6 female; Kur- undia, 23 .111.20 12, 1 male; Jeeragaon, 26.III.2012, 1 male and 1 female; Hatguda, 29.III.2012, 1 male; Taraguda, 16.IV.20 12, 3 males and 3 females; Kalcha, 24.IV.20 12, 1 male; Bhatiguda, 2.VI.2012, 1 male; Machkote Range, 7.VI.2012, 1 female; Rawanapat, 23.X.2013, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily TERATODINAE Genus Teratodes Brulle, 1835 28. Teratodes monticollis (Gray, 1832) Gryllus monticollis - Gray, 1832: 215 Teratodes monticollis - Shishodia, 2000: 52 Examined material. Chhattisgarh; Bastar, Jag- dalpur range, 29.VIII.2011, 2 females. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Superfamily PYRGOMORPHOIDEA Family PYRGOMORPHIDAE Genus Atractomorpha Saussure, 1862 29 . Atractomorpha crenulata (Fabricius, 1793) Truxalis crenulata - Fabricius, 1793: 28 Atractomorpha crenulata - Shishodia, 2000: 42 Examined material. Chhattisgarh; Bastar, Mal- gaon, 23. XI. 2011,1 male; 28.VII.2011, 1 female; Dagania, 29.VIII.2011, 1 female; Sonarpara Beat, 17.X. 2011, 1 male; Makdi range, 10.XI.2011, 1 male; Neganar Village, 4.1.2012, 1 male; Nathguda Village, 24.1.2012, 1 male; Taraguda, 12.III.2012, 1 male; Ulnar Village, 16.III.2012, 1 male; Kurundi, 23. 111. 2012. 1 male; Hatguda, 29.III.2012, 1 female; Machkot range, 7.VI.2012, 2 males; Bhatiguda Vil- lage, 18.VII.2012, 1 male; Pushpal, 1.VIII.2013, 2 males. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Aularches Stal, 1873 Systematic account of Orthoptera fauna of Bastar district, Chhattisgarh, India 45 30. Autarches miliaris miliaris (Linnaeus, 175 8) (*) Gryllus ( Locusta ) miliaris - Linnaeus, 1758: 432 Aularches miliaris miliaris - Mandal & Yadav, 2007: 190 Examined material. Chhattisgarh; Bastar, Erikpal Village, 16.VIII.2011, 1 female; Kanker, 27.VII.2011, 1 female; 28.VII.2011, 1 female; Jag- dalpur city, 29.VIII.2011, 1 female; Jagdlapur range, 30. VIII. 2011, 1 female; Asna Village, 2. 11. 2012, 1 female; Malegaon, 10.III.2012, 1 fe- male; Jhiriya Bahara, 20.III.2012, 1 female. Distribution in Chhattisgarh. Bastar. Remark. New record from Chhattisgarh State. Genus Chrotogonus Audinet-Serville, 1838 31. Chrotogonus ( Chrotogonus ) trachypterus trachypterus (Blanchard, 1836) Ommexycha trachypterus - Blanchard, 1836: 618 Chrotogonus (C.) trachypterus trachypterus - Shishodia, 2000: 40 Examined material. Chhattisgarh; Bastar, Amaguda, 24.VIII.2011, 1 male and 1 female; Bhanpur, 19.X.2011, 2 females; 21.X.2011, 1 male and 1 female; Pipalvond Beat, 22.X.2011, 1 male and 1 female; Malegaon, 23.XI.2011, 1 female; Bel- guda Village, 16.1.2012, 3 females; Natguda Vil- lage, 24.1.2012 2 males and 3 females; Neganar Village, 4.1.2012, 1 male; Malegaon, 10.III.2012, 1 female; Kohkapal, 14.III.2012, 1 male and 5 fe- males; Hathguda, 29.III.2012, 1 female; Taraguda, 16.IV.2012, 1 female; Kalcha, 24.IV.2012, 2 males; Ulnar, 22. XI. 2011, 1 male and 1 female. Distribution in Chhattisgarh. Bstar, Bilaspur, Kabirdham, Raipur. Genus Poekilocerus Audinet-Serville, 1831 32 . Poekilocerus pictus (Fabricius, 1775) Gryllus pictus - Fabricius, 1775: 289 Poekilocerus pictus - Kirby, 1914: 172 Examined material. Chhattisgarh; Bastar, Mo- grapol Village, 7.1.2012, 1 female; Nakaguda, 19.1.2012, 1 male; Amaguda, 2.III.2012, 1 male. Distribution in Chhattisgarh. Bastar and Raipur. Superfamily TETRIGOIDEA Family TETRIGIDAE Subfamily SCELIMENINAE Genus Criotettix Bolivar, 1887 33. Criotettix bispinosus (Dalman, 1818) Acrydium bispinosum - Dalman, 1818: 77 Criotettix bispinosus - Bolivar, 1887: 185, 223, 226 Criotettix bispinosus - Gunther, 1938: 134 Examined material. Chhattisgarh; Bastar, Sonarpal Beat, 17.X.2011, 1 female. Distribution in Chhattisgarh. Bastar. Genus Euscelimena Gunther, 1938 34 . Euscelimena harpago (Audinet-Serville, 1839) Tetrix harpago - Audinet-Serville, 1839: 763 Euscelimena harpago - Hebard, 1929: 572 Examined material. Chhattisgarh; Bastar, Na- kaguda, 19.1.2012, 1 male; Asna Village, 2.II.2012, 2 females; 4.II.2012, 1 female; Malgaon, 10.III.2012, 2 females; Kohkapal, 14.III.2012, 2 females; Machkote Range, 7.VI.2012, 1 male and 1 female; Amaguda, 2.III.2012, 1 male; Kurundi, 23.III.2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily TETRIG1N AE Genus Ergatettix Kirby, 1914 35. Ergatettix dorsiferus (Walker, 1871) Tettix dorsifera - Walker, 1871: 825 Ergatettix dorsifera - Shishodia, 1999: 42 Examined material. Chhattisgarh; Bastar, Makdi, 9.XI.201, 1 male; Nakaguda, 19.1.2012, 1 male; Amaguda, 2.III.2012, 1 male; Malgaon, 9. III. 2012, 1 male; Ulner Village, 16.III.2012, 1 male. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Hedotettix Bolivar, 1887 36. Hedotettix gracilis (de Haan, 1842) Acridium {Tetrix) gracile - de Haan, 1842: 167-169 Hedotettix gracilis - Shishodia, 2000: 36 46 Sunil Kumar Gupta Examined material. Chhattisgarh; Bastar, Jag- dalpur range, 28. VII. 2011, 2 males and 1 female; Sonarpali Beat, 17.X.20011, 2 males. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Infraorder TRIDACTYLIDEA Superfamily TRIDACTYLOIDEA Family TRIDACTYLIDAE Brunner, 1882 Subfamily TRIDACTYLINAE Genus Tridactylus Olivier, 1789 37. Tridactylus thoracicus Guerin, 1844 Tridactylus thoracicus - Guerin, 1844: 336 Tridactylus thoracicus - Shishodia & Tandon, 1987: 128 Examined material. Chhattisgarh; Bastar, Neganar Village, 5.1.2012, 1 male, DC; 31.1.2012, 1 female, DC; Amaguda, 2.III.2012, 1 female, DC; Gariya bahar river, 24.III.2012, 1 male, DC; Mon- grapal Village, 1.1.2012, 1 male, DC; 7.1.2012, 1 fe- male, DC. Distribution in Chhattisgarh. Bilaspur and Raipur. Suborder ENSIFERA Infraorder OEDISCHIOIDEA Superfamily GRYLLOIDEA Family Gryllidae Subfamily Gryllinae Genus Loxohlemmus Saussure, 1877 38. Loxoblemmus haani Saussure, 1877 (*) Loxohlemmus haani - Saussure, 1877: 257 Loxohlemmus haani - Vasanth, 1993: 46 Examined material. Chhattisgarh; Bastar, Nandpurabeat, 20.X.2011, 1 male. Distribution in Chhattisgarh. Basatr, Bilaspur. Remark. New record from Chhattisgarh State. Genus Modicogryllus Chopard, 1961 Subgenus Modicogryl l us Chopard, 1961 39 . Modicogryllus (Modicogryllus) confirma- tus (Walker, 1859) Acheta confirmata - Walker, 1859: 221 Modicogryllus confirmatus - Tandon et al., 1976: 170 Examined material. Chhattisgarh; Bastar, Nandpura Beat, 20.X.2011, 1 female; Jagdalpur range, 11. XI. 2011, 1 female; Gariya bahar river, 20. 111. 2012, 1 female; Malgaon, 10.III.2012, 1 fe- male. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Phonarellus (Gorochov, 1983) Subgenus Phonarellus Gorochov, 1983 40. Phonarellus ( Phonarellus ) minor Chopard, 1959 Gymnogryllus minor - Chopard, 1959: 1 Phonarellus ( Phonarellus ) minor - Gorochov, 1983: 323 Examined material. Chhattisgarh; Bastar, Bhanpuri, 20.X.2011, 1 female; Mongrapal Village, 7.1.2012, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Teleogryllus Chopard, 1961 Subgenus Macroteleogryllus Gorochov, 1988 41. Teleogryllus {Macroteleogryllus) mitratus (Burmeister, 1838) Gryllus mitratus - Burmeister, 1838: 734 Teleogryllus mitratus - Gupta et al., 2008: 120 Examined material. Chhattisgarh; Bastar, Jag- dalpur range, 24.VIII.2011, 1 female. Distribution in Chhattisgarh. Bastar and Bilaspur. Subfamily NEMOBIINAE Genus Paranemohius Saussure, 1877 42. Paranemobius pictus (Saussure, 1877) Pseudonemobius pictus - Saussure, 1877: 67 Paranemohius pictus - Shishodia, 2000: 70 Examined material. Chhattisgarh; Bastar, Neganar Village, 4.1.2012, 1 female. Distribution in Chhattisgarh. Bastar. Subfamily OECANTHINAE Genus Oecanthus Audinet-Serville, 1831 Systematic account of Orthoptera fauna of Bastar district, Chhattisgarh, India 47 43. Oecanthus indicus Saussure, 1878 Oecanthus indicus - Saussure, 1878: 454. Oecanthus indicus - Shishodia, 2000: 71 Examined material. Chhattisgarh; Bastar, Ericpal Village, 24.11.2012, 1 female. Distribution in Chhattisgarh. Bastar and Bilaspur. Family TRIGONIDIIDAE Genus Anaxipha Saussure, 1874 44. Anaxipha sp. Anaxipha - Saussure, 1874: 370 Anaxipha - Vasanth, 1993: 108 Examined material. Chhattisgarh; Bsatar, Jag- dalpur Forest, 15.VII.2011, 1 male and 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Trigonidium Rambur, 1839 Subgenus Trigonidium Rambur, 1839 45. Trigonidium ( Trigonidium ) cicindeloides Rambur, 1839 Trigonidium cicindeloides - Rambur, 1839: 39 Trigonidium cicindeloides - Shishodia, 2000: 74 Examined material. Chhattisgarh; Bastar, Bhanpuri, 19.X.2011, 1 female. Distribution in Chhattisgarh. Bastar, Raipur. Family GRYLLOTALPIDAE Genus Gryllotalpa Latreille, 1802 46. Gryllotalpa africana Beauvois, 1805 Gryllotalpa africana - Palisot de Beauvois, 1 805 : 229 Gryllotalpa africana - Shishodia, 2000: 64 Examined material. Chhattisgarh; Bastar, Nand- pura Village, 20.X.2011, 1 male; Bhanpur, 21.X.2011, 1 male; Rampal, 19.i.2012, 1 female; Hathguda, 29.III.2012, 1 male. Distribution in Chhattisgarh. Bastar, Bila- spur, Raipur. Superfamily TETTIGONIOIDEA Family TETTIGONIIDAE Subfamily CONOCEPEtALINAE Genus Conocephalus Thunberg, 1815 Subgenus An isoptera Latreille, 1829 47. Conocephalus ( Anisoptera ) maculatus (Le Guillou, 1841) Xiphidion maculatum - Le Guillou, 1841: 294 Conocephalus maculatus - Chandra et al., 2007: 2684 Examined material. Chhattisgarh; Bastar, Jag- dalpur Forest, 15. VII. 2011, 1 female; Bhanpuri Forest, 20.X.2011, 1 female; Mangrapal Village, 7.1.2012, 2 females; Belguda Village, 16.1.2012, 1 female; 18.1.2012, 1 female; Naganar Village, 31.1.2012, 1 male; Asna Village, 4. II. 2012, 1 fe- male; Dongaghat, 8.II.2012, 1 female; Amaguda Village, 2. III. 2012, 1 male; Malegaon, 10.III.2012, 1 female; Jagdalpur, 12.III.2012, 1 male; Taraguda, 12. 111. 2012, 1 male; Bhanpuri, 15.III.2012, 1 fe- male; Bhatiguda Village, 2.VI.2012, 1 male. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily MECOPODINAE Genus Mecopoda Audinet-Serville, 1831 48. Mecopoda elongata elongata (Linnaeus, 1758) Gryllus ( Tettigonia ) elongatus - Linnaeus, 1758: 429 Mecopoda elongata - Barman, 2003: 195 Examined material. Chhattisgarh; Bastar, Taraguda, 16.IV.20 12, 1 female. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily PHANEROPTERINAE Genus Elimaea Stal, 1874 Subgenus Orthelimaea Karny, 1926 49. Elimaea ( Orthelimaea ) securigera Brunner von Wattenwyl, 1878 Elimaea ( Orthelimaea ) securigera - Brunner von Wattenwyl, 1878: 93 Elimaea ( Orthelimaea ) securigera - Barman, 2000: 264 Examined material. Chhattisgarh; Bastar, Kotamsur, 27.VII.2011, 1 male. 48 Sunil Kumar Gupta Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Genus Himertula Uvarov, 1 940 50 . Himertula kinneari (Uvarov, 1923) Himerta kinneari - Uvarov, 1923: 661 Himertula kinneari - Ingrisch & Shishodia, 2000: 20 Examined material. Chhattisgarh; Bastar, Jagdalpur range, 30.VIII.2011, 1 female. Distribution in Chhattisgarh. Bastar and Raipur. Genus Phaneroptera Audinet-Serville, 1831 Subgenus Phaneroptera Audinet-Serville, 1831 51. Phaneroptera gracilis Burmeister, 1838 Phaneroptera gracilis - Burmeister, 1838: 690 Phaneroptera gracilis - Shishodia, 1999: 36 Examined material. Chhattisgarh; Bastar, Ericpal, 24.11.2012, 1 male. Distribution in Chhattisgarh. Bastar, Bilaspur, Raipur. Subfamily Pseudophyllinae Genus Sathrophyllia Stal, 1874 52. Sathrophyllia rugosa (Linnaeus, 1758) Gry llus ( Tettignonia ) rugosa - Linnaeus, 1758: 430 Sathrophyllia rugosa - Beier, 1962: 199-200 Examined material. Chhattisgarh; Bastar, Erikpal, 22.VII.2011, 1 female. Distribution in Chhattisgarh. Bastar, Raipur. ACKNOWLEDGEMENTS The author is grateful to Dr. Kailash Chandra, Director, Zoological Survey of India (Kolkata) for providing necessary facilities and encourage- ments. Thanks are also due to CAMPA (Compens- atory Afforestation Fund Management and Planning Authority) for funding the project, and to Chhattisgarh Forest Department for providing necessary permissions and support to carry out the present work. REFERENCES Audinet-Serville J.G., 1831. Revue methodique des Orthopteres. Annales des Sciences Naturelles, 22: 28-65, 134-162,262-292. Audinet-Serville M., 1839. Histoire naturelle des insects: Orthopteres. 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Catalogue of the specimens of Dermaptera Saltatoria in the collection of the British Museum, Part V, Supplement to the Catalogue of Dermaptera Saltatoria. British Museum, London, 4: 4 + 811-850 + 43 + 116. Willemse C., 1951. Overdruk uit .publicities van Het Natuurhistorisch Genoots chap in Limberg Vol. IV, 15th August, 1951. Synopsis of the Acridoidea of the Indo Malayan and acijacent regions (Insecta: Orthoptera). Biodiversity Journal, 2016, 7 (1): 51-54 First record of Brachytron pratense (Miiller, 1764) in Sicily (Odonata Aeshnidae) Paolo Galasso 1 *, Nadia Curcuraci 1 & Alessandro Marietta 2 'Stiftung Pro Artenvielfalt®, MeisenstraBe 65, 33607 Bielefeld, Germany; email: paolo_galasso@hotmail.com, nadiacurcuraci@yahoo . it department of Biological, Geological and Environmental Sciences - section of Animal Biology “M. La Greca”. University of Catania, via Androne 81, 95124 Catania, Italy; email: amarlet@unict.it ^Corresponding autor ABSTRACT Brachytron pratense (Muller, 1764) is a small Odonata Aeshnidae widespread throughout most of Europe and Central-northern Italy, but up to now never recorded in Sicily. During the spring 2015, some specimens of this species were observed and photographed for the first time at the swamp lake “Pantano Cuba”, in the southeast coast of Sicily, near to Pachino (Syracuse). This record represents now the southernmost Italian locality for this species. KEY WORDS Pantano Cuba; Odonata; dragonflies; Stiftung Pro Artenvielfalt; Sicily. Received 08.03.2016; accepted 22.03.2016; printed 30.03.2016 INTRODUCTION Brachytron pratense (Muller, 1764) is a small Odonata Aeshnidae that is often confused with others species belonging to the genus Aeshna Vander Linden, 1820; however, unlike these, it can be observed in flight early in March and it presents some peculiar morphological characters. It is a generally localised species, with a Central European distribution which extends to Balkan and Mediter- ranean region. Its range includes the west of the Urals, France (Corsica included), Netherlands, Ireland, United Kingdom, Switzerland, Austria, Germany, Slovenia, Croatia, Czech Republic, Slovakia, Greece, Denmark, Finland, Norway, Sweden, Poland, Romania, Estonia, Fatvia, Fithuania, Belarus and Russia (Askew 2004; Dijkstra & Fewington, 2006). In Italy it is an uncommon species and appears more widespread in northern regions, with the exception of Figuria and Val D' Aosta (Fig. 1). However, in the central and southern regions only few isolated localities are known, so that a good definition of areal borders is precluded (Riservato et al., 2014a). Until now the species had never been reported for Sicily (Riservato et al., 2014b) and the known southernmost record was in Calabria, near Famezia Terme (Fig. 1). Therefore, this new record extends southward the known Italian distribution of this species and represents now its southernmost Italian locality. MATERIAL AND METHODS During a biodiversity monitoring program promoted by the German "Stiftung Pro Artenvielfalt - Pro Biodiversity Foundation" at the swamp lake “Pantano Cuba”, since April 2015 we have ob- served and photographed some specimens of B. 52 Paolo Galasso et alii pratense. Data were collected during odonatologic surveys from March 2015 to December 2015. Sur- veys have been conducted regularly every week at the same location and with the same method: tran- sects traversed on foot, collecting and releasing the specimens with aerial nets for identification. Moreover several macrophotos have been made on- site using a digital SLR camera. The species shows characters so unmistakable that it was not necessary to kill and preserve the Figure 1. Distribution map of Brachytron pratense in Italy. Red arrow shows the new record area (edit from CKmap). Figure 2. Location of "Riserva dei Pantani della Sicilia sud-orientale" (Pachino, Syracuse), new locality record for Brachytron pratense (from Google Earth). specimens captured. So they were released imme- diately after the identification. The place of occurrence, Pantano Cuba (36°42’26.71”N; 15°1 , 39.15 ,, E), along a complex of others 7 swamp lakes with different sizes, con- stitute a very important coastal wetland which was part of a natural reserve named "Riserva dei Pantani della Sicilia Sud-orientale" (Fig. 2), whose estab- lishment was cancelled on May 2015. The swamp, which is located less than 500 meters from the sea, lies entirely in the municipality of Pachino, in the province of Syracuse; it has an extension of 63 hec- tares and it is characterized by brackish and still wa- ters with abundant aquatic vegetation represented mainly by Ruppia maritima L., vegetation helo- phytic with Phragmites australis (Cav.), Bol- boschoenus maritimus (L.) Palla, Juncus acutus L., Juncus maritimus Lam. and Tamarix africana Poir., as well as by halophytic vegetation zones with Arth- rocnemum fruticosum (L.) and Inula crithmoides L. Near to the swamp shores there are also idle land, now entirely covered by grassy vegetation and several trees of Acacia saligna Labill. RESULTS Brachytron pratense adults have a length of 54- 63 mm and a wingspan of 68-74 mm. They are unmistakable, characterized by hairy thorax and abdomen, densely covered by thin setae (Figs. 3- 6). The sides of thorax are green, distinctly inter- rupted by two complete black lines (Fig. 3). The wings with a narrow and elongated pterostigma (Fig. 4). Males abdomen black and cylindrical, not narrowed at the base, with pairs of elongated blue spots on almost all segments and a diagnostic central yellow dot on the first abdominal tergite S 1 (Figs. 4, 6). The females (Figs. 3, 5) are similar to males, except for abdomen stout, browner with greenish-yellow (not blue) spots (Askew, 2004; Dijkstra & Fewington, 2006). During the surveys at Pantano Cuba, several specimens of B. pratense were observed in at least four different occasions, always in the same site; they were adults of both sexes: - April 9, 2015, 1 female (Figs. 3, 5): it was caught near one of the fallow fields, about 60 meters from the main water body; it was photo- graphed and released. First record of Brachytron pratense (Muller, I 764) in Sicily (Odonata Aeshnidae) 53 Figures 3. Brachytron pratense female (Pachino, Pantano Cuba; 9.IV.2015): in hand (ventral-lateral view), showing the typical hairy body. Figure 5. Dorsal view of the same specimen. Figure 4. Brachytron pratense male (Pachino, Pantano Cuba; 1.V.2015): dorsal view (pt, pterostigma). Figure 6. Brachytron pratense male (Pachino, Pantano Cuba; 23.IV.2015): dorsal-lateral view (Photos by P. Galasso). -April 23, 2015, 1 male (Fig. 6): it was observed and photographed on a branch of Acacia saligna near a small ditch about 130 meters from the main water body. - May 1, 2015, 2 males: they showed territorial behaviour, one of them was photographed (Fig. 4); they were observed in a wet meadow of Inula crithmoides a few meters from the main water body. - May 7, 2015, 1 male (not photographed): it was observed in full predatory activities through open meadows about 100 meters from the main water body. CONCLUSIONS These records add an important and valuable contribution to the Italian and European odonato- logy and especially to the study of B. pratense dis- tribution and ecology. 54 Paolo Galasso et alii Pantano Cuba is the first Sicilian site for this species and the southernmost of Italy and Europe; it also highlights the undoubted importance of research projects and monitoring of high conserva- tion value areas such as the Pantano Cuba, often underestimated and not subject to the strict retention policies and management of biodiversity which they would deserve. ACKNOWLEDGEMENTS We wish to thank “ODONATA.IT - Societa Italiana per lo Studio e la Conservazione delle Libellule” (Carmagnola, Torino, Italy), for further confirmation of identification through the analysis of the photographic material. REFERENCES Askew R.R., 2004. The dragonflies of Europe. Harley Books. Colchester, 308 pp. Dijkstra K.D. & Lewington R., 2006. Field Guide to the Dragonflies of Britain and Europe. British Wildlife Publishing, Gillingham, 320 pp. Riservato E., Fabbri R., Festi A., Grieco C., Hardersen S., Landi F., Utzeri C., Rondinini C., Battiston A. & Teofili C. (compilatori) 2014a. Fista Rossa IUCN delle libellule Italiane. Comitato Italiano IUCN e Ministero dell’ Ambiente e della Tutela del Territorio e del Mare, Roma, 39 pp. Riservato E., Festi A., Fabbri R., Grieco C., Hardersen S., Fa Porta G., Fandi F., Siesa M.E. & Utzeri C., 2014b. Odonata - Atlante delle libellule italiane - preliminare. Societa Italiana per lo Studio e la Conservazione delle Fibellule - Edizioni Belvedere, Fatina, “le scienze” (17), 224 pp. Biodiversity Journal, 2016, 7 (1): 55-57 First record of Callistochiton pachylasmae (Monterosato, 1 879) for the Adriatic Sea (Polyplacophora Callistoplacidae) Bruno Amati 1 & Marco Oliverio 2 'Largo Giuseppe Veratti 37/D, 00146 Rome, Italy; e-mail: bnmo_amati@yahoo.it 2 Dipartimento di Biologia e Biotecnologie ‘Charles Darwin’, Sapienza Universita di Roma, Viale dell’Universita 32, 00185 Rome, Italy; e-mail: marco.oliverio@uniromal.it ABSTRACT It is reported the first record of Callistochiton pachylasmae (Monterosato, 1879) for the Adriatic Sea. It a very rare and peculiar polyplacophoran species (Callistoplacidae Pilsbry, 1893). Actually, The few known records span a wide Mediterranean range and extend to the neighbouring Atlantic. KEY WORDS Callistochiton pachylasmae', polyplacophoran species; first record; Adriatic Sea. Received 17.01.2016; accepted 09.02.2016; printed 30.03.2016 INTRODUCTION Callistochiton pachylasmae (Monterosato, 1879) (original combination: Chiton pachylasmae Monterosato, 1879 ex Seguenza G. ms) is a very peculiar polyplacophoran species ( Callistoplacidae Pilsbry, 1893: Bouchet et al., 2016; Gofas & Le Renard, 2016), easily recognizable by its peculiar sculpture, in particular for the presence of 7 radial ridges on the cephalic plate. Its distinctiveness, along with its apparently isolated fossil history in Europe, traced back to at least the Pleistocene (Dell’ Angelo et al., 1998), brougth Dell’ Angelo & Oliverio (1997) to allocate it in a subgenus on its own: Allerychiton Dell’ Angelo et Oliverio, 1997. DISCUSSION AND CONCLUSION Callistochiton pachylasmae is a rare species, and it has been treated seldom in the literature (Monterosato, 1879; Sabelli, 1971; Ferreira, 1979; Kaas, 1981; van Belle, 1983, 1988; Gaglini, 1985; Pizzini & Oliverio, 1993; Giovine & Dell’ Angelo, 1993; Kaas & van Belle, 1994; Dell’ Angelo & Oliverio, 1997; Dell’Angelo et al., 1998; Anto- niadou et al., 2005; Koulcouras, 2010). The few known records span a wide Mediterranean range and extend to the neighboring Atlantic (Fig. 1). It is noteworthy that the generic record from Spain in the Iberian Fauna Databank (Ramos, 2010) could not be linked to an actual, published record (J. Templado, pers. comm.) and therefore could not be plotted in the map (Fig. 1). However, despite the wide range, there was so far remarkable lack of findings in the Adriatic Sea. The present record consist of a single cephalic plate, 0.82 x 1.37 mm (Fig. 2), retrieved by sorting a sample of bioclastic sediment with limited or- ganogenous component, collected by SCUBA diving at Lastovo Island (Croatia), 38 m depth (Alessandro Raveggi, Florence, legit). This is the first record from the Adriatic Sea, and represents the northernmost known record for the species. 56 Bruno Amati & Marco Oliverio Figure 1. Known records of Callistochiton pachylasmae (Monterosato, 1879). 1) W of Cape Yubi, Morocco, -500 m, 1 spe- cimen now lost (Kaas, 1981). 2) Punta Longa “Secca Galera”, Favignana Island -33 m, 1 cephalic plate (Dell’ Angelo & Oliverio, 1997). 3) Pantelleria Island, -53.4 m (Pizzini & Oliverio, 1993). 4) Strait of Messina, coralligenous, 1 specimen (holotype: Monterosato, 1879). 5) S. Maria di Catanzaro, Pleistocene, 1 cephalic plate (Dell’ Angelo et al., 1998). 6) Lastovo Island (Croatia), -38 m (this work). 7) Kelyfos Island, -30 m, 1 specimen (Antoniadou et al., 2005; Koukouras, 2010). 8) Ormos Panagias -35/40 m, Sithonia, 1 intermediate plate (Dell’ Angelo & Oliverio, 1997). Figure 2. Callistochiton pachylasmae (Monterosato, 1879). Lastovo Island (Croatia), 38 m depth. Cephalic plate, height 0.82, width 1.37 mm. REFERENCES Antoniadou C., Koutsoubas D. &. Chintiroglou C.C., 2005. Mollusca fauna from infralittoral hard substrate assemblages in the North Aegean Sea. Belgian Journal of Zoology, 135: 119-126. Belle R.A. van, 1983. The Systematic Classification of the Chitons. Information de la Societe Beige de Malacologie, Ser. 11 (1-3): 1-178. Belle R.A. van, 1988. De Europese keverslakken (Poly- placophora) [The European Polyplacophora], Vita Marina Jan.-Mrt: 9-132 (look up in IMIS). Bouchet P., Marshall B., Schwabe E. & Gofas S., 2016. Callistochiton Dali, 1879. In: MolluscaBase (2016). Accessed through: World Register of Marine Species athttp://www.marinespecies.org/aphia.php?p= taxde- tails&id=138084 on 2016-01-04 Dell’Angelo B. & Oliverio M., 1997. A new poly- placophoran subgenus for the Northeast Atlantic (Ischnochitonidae: Callistoplacinae). Bollettino Ma- lacologico, 32: 145-150. Dell’Angelo B., Vazzana A. & Bertolaso L., 1998. Ritro- vamento di piastre fossili di Callistochiton (Mollusca: Polyplacophora) nel Plio-Pleistocene della Calabria. Bollettino Malacologico, 33 (1997): 139-140. Ferreira A. J., 1979. The Genus Callistochiton Dali, 1879 (Mollusca: Polyplacophora) in the Eastern Pacific, with the Description of a New Species. The Veliger, 21; 444-466. Gaglini A.,1985. Classe Anphineura. In: Settepassi F., Atlante Malacologico. I Molluschi Marini viventi nel Mediterraneo. Roma, Vol. III., 19 pp., 13 tav. Giovine F. & Dell’Angelo B., 1993. Elenco dei Mol- luschi rinvenuti nello Stretto di Messina. Poly- placophora. Lavori S.I.M. 24, III Congresso S.I.M., Parma 1990: 157-170. Gofas S. & Le Renard J. (Eds.), 2016. CLEMAM: Check List of European Marine Mollusca. Available from First record of Callistochiton pachylasmae for the Adriatic Sea (Polyplacophora Callistoplacidae) 57 http://www.somah.asso.fr/clemam/biotaxis.php7XM 0251&header=l on 2016.01.02. Kaas P., 1981. Chitons (Mollusca, Polyplacophora) pro- cured by the CANCAP I- VII expeditions, 1976- 1986. Zoologische Mededelingen, 65: 89-98. Kaas P. & van Belle R.A., 1994. Monograph of living chitons. Vol. 5, Additions to Volumes 1-4. Koukouras A., 2010. Checklist of marine species from Greece. Aristotle University of Thessaloniki. Assembled in the framework of the EU FP7 PESI project. Monterosato T.A. di Maria di, 1879. Enumerazione e sinon- imia delle conchiglie mediterranee. Parte I. Mono- grafia dei Chitonidi del Mediterraneo. Giornale di Scienze Naturali ed Economiche di Palermo, 14: 1-31. Pizzini M. & Oliverio M., 1993. Ritrovamento di Cal- listochiton pachylasmae (Monterosato, 1878 ex Seguenza G. ms) in Mediterraneo (Polyplacophora, Ischnochitonidae). Notiziario C.I.S.Ma, 14 (1992): 41-42. Ramos M., 2010. IBERFAUNA. The Iberian Fauna Da- tabank. Sabelli B., 1971. Revisione del Chiton pachylasmae Monterosato. Atti XL Convegno U.Z.I., Bollettino Zoologico, 38: 561. Biodiversity Journal, 2016, 7 (1): 59-66 Monograph Jujubinus errinae n. sp. (Gastropoda rochidae from the Strait of Messina, Mediterranean Sea Carlo Smriglio 1 , Paolo Mariottini 1 * & Salvatore Giacobbe 2 'Dipartimento di Scienze, Universita “Roma Tre”, Viale Marconi, 446, 00146 Roma, Italy; e-mail: csmriglio@alice.it; paolo. mariottini@uniroma3 . it 2 Dipartimento di Scienze Biologiche e Ambientali, Universita di Messina, Viale Stagno D'Alcontres, 98166 Messina, Italy; e-mail: sgiacobbe@unime.it Corresponding author, email: paolo.mariottini@uniroma3.it ABSTRACT A new species of the gastropod family Trochidae, Jujubinus errinae n. sp., from the Mediter- ranean Sea is described based on shell characters. The new taxon was compared with the most closely related species showing marked sculpture and from relatively deep water habitat, J. catenatus Ardovini, 2006, J. montagui (Wood, 1828) and J. tumidulus (Aradas, 1846). The species, which is known from the type locality only, the Strait of Messina, might be strictly associated to the endemic hydrocoral Errina aspera (Linnaeus, 1767) beds (Hydrozoa Stylas- teridae). KEY WORDS Trochidae; Recent; Jujubinus errinae', new species; Mediterranean Sea. Received 30.10.2015; accepted 16.01.2016; printed 30.03.2016 Proceedings of the Ninth Malacological Pontine Meeting, October 3rd-4th, 2015 - San Felice Circeo, Italy INTRODUCTION The Strait of Messina has been considered a separate Mediterranean biogeographic microsector inhabited by rich benthic communities and some peculiar assemblages that are unknown in other Mediterranean regions (Bianchi, 2004). From this specific area, a survey of the species of the genus Jujubinus Monterosato, 1884 (Gastropoda Trochidae) has been carried out on samples from hard and soft circalittoral bottoms, which revealed the presence of trochidae shells not recognizable as a known species. The specimens, once compared with Juju- binus catenatus Ardovini, 2006, J. montagui (Wood, 1828) and J. tumidulus (Aradas, 1846), the most closely related species showing marked sculp- ture and from relatively deep water habitat, were attributed to a new species of this genus, J. errinae n. sp., which is here described. ACRONYMS. The materials used for this study are deposited in the following private and Museum collections: Carlo Smriglio and Paolo Mariottini, Rome, Italy (CS-PM); DiSBA Benthic Ecology laboratory Messina, Italy (DiSBA); Giuseppe Notaristefano, Messina, Italy (GN); Museo Civico di Zoologia, Rome, Italy (MCZR); Museo di Zoologia Bologna, Bologna, Italy (MZB); Renato Marconcini, Reggio Calabria, Italy (RM); Walter Renda collection, Reggio Calabria, Italy (WR); Bruno Amati, Roma, Italy (BA); Ermanno Quaggiotto, Logare, Vicenza, Italy (EQ). Other acronyms used in the text: Height (H); Interdepar- tmental Laboratory of Electron Microscopy, Rome, Italy (LIME); Monterosato (MTS); Scanning Elec- tron Microscopy (SEM); specimens (sps); station (st); Width (W). 60 Carlo Smriglio etalii MATERIAL AND METHODS Hard and soft bottom samples containing the new species were collected in the Strait of Messina, central Mediterranean, in several cruises sponsored by the University of Messina. In particular, dredging was carried out during the “POP 95” cruise (13 to 31 July 2015), at 100 m depth (DG04: 38°14'45" N, 15°37'36" E), and arising fromll5 m to 90 m, along a steep rocky floor (DG001: 38°14'45" N, 15°37'28" E). During the same cruise, van Veen grab samples were collected in Rada Paradiso [Station (St) 02: 38°13'27"N, 15°36'02"E], 201 m depth. A further grab sample, collected by the R/V Coopernaut Franca in the framework of the POR-CAL 2008 project, was carried out in October 2008, on the slope of the Gioia Basin (St IB: 38°18'6941N, 15°45'5710E), 371 m depth. Bioclastic sediment samples were also collected during SCUBA diving on the bottoms of the Strait of Messina, at a depth of 40-50 m (38°15 , 36”N, 15°43 , 08”E). Sediment samples were sieved through a 1 mm mesh and the residue was sorted using a stereomicroscope. Among the sorted material, shells of an undescribed species of Jujubinus, represented by 21 sps, to- gether juveniles and fragments not included in the type series, were separated and described herein as J. errinae n. sp. Additional material examined from CS-PM collection: 3 sps of J. catenatus from the Sicily Channel, estimated depth 90 m; about 100 sps of J. montagui from Anzio, Central Tyrrhenian Sea, 50 m; 9 sps from Sfax, Tunisia, 100 m; over 200 sps of J. tumidulus from Lampedusa Island, Sicily Channel, dredged by fishing boats, estimated depth 70-80 m; 11 sps from Linosa Island (Punta Calcarello), Sicily Channel, 36 m. Current system- atics is based on WoRMS (Gofas & Bouchet, 2015). Scanning Electron Microscopy (SEM) photo- graphs were taken at the Interdepartmental Laborat- ory of Electron Microscopy (LIME, Universita “Roma Tre”, Rome, Italy), using a Philips XL30. SYSTEMATICS Classis GASTROPODA Cuvier, 1795 Familia TROCHIDAE Rafmesque, 1815 Genus Jujubinus Monterosato, 1884 Type species (by subsequent designation of Crosse, 1885) Trochus matoni Payraudeau, 1826 Jujubinus errinae n. sp. (Figs. 1-18,37) Diagnosis. Small and slightly turriculate shell; sculpture of incised spiral lines; strong prosocline lamellae between spiral cords. Examined material. The holotype (MZB60155) and paratypes A-H (DiSBA) from the type loc- ality: Strait of Messina, (38°14'45”N 15°37'36”E), Sicily, Mediterranean Sea, dredging DG04, 100 m depth; paratypes I (DiSBA) from 38°14'45" N, 15°37'28" E, dredging DG001, St 5, 90-115 m; paratype L (DiSBA) from 38 0 13'27’’ N, 15°36'02" E, dredging PIC02, Rada Paradiso, St 2, 90-115 m; paratype M (WR) from 38°14'45" N, 15°37'28" E, dredging DG001, St 5, 90-115 m; paratype N-R (CS-PM) from 38°15'36" N, 15°43'08" E, 40-50 m depth; paratypes S-U (GN); paratype V (RM) from 38°15'36" N, 15°43'08" E, 40-50 m depth; paratype X (BA) from 38°15'36" N, 15°43'08" E, 44 m depth; paratype Y (EQ), from 38°15'36" N, 15°43'08" E, 40-50 m depth. Description of holotype. Shell of relatively small size for the genus, height (H) 4.9 mm, width (W) 4.0 mm, conical, slightly shiny. Protoconch about 1.5 whorls, smooth, with a diameter of 280 pm. Teleoconch of 4.5 slightly convex whorls. Sculpture of 6 closely set abapical spiral cords of about the same strength, strongly carved by strong tubercles, including the 2 peripheral ones forming the basal cord, and 6 regularly spaced, basal spiral cords narrow and well engraved, with very evident lamellae in the interspaces. First two whorls of the teleoconch showing the basal cord strongly rippled, remaining teleoconch whorls with a flat basal cord. Suture incised. Teleoconch surface covered by barely visible prosocline growth striae, irregularly set. Base convex, umbilicus closed and covered with a white callus. Aperture quadrangular, with the columellar callus thickened in the middle portion and internally whitish nacreous. Colour of proto- conch whitish, teleoconch reddish-creamy, with red spiral cords interrupted by short white spots. The same chromatic pattern is shown by the basal cords. Animal unknown. A new Jujubinus from the Mediterranean Sea 61 Variability. Shell H ranging from 4.5 to 6.0 mm and W from 3.8 to 4.7 mm. Protoconch dia- meter from 260 to 290 pm. Teleoconch varying from 4 to 4.5 whorls. Spiral and basal cords both ranging from 5 to 6, according to the H of the shell (6 in adult specimens). Umbilicus is closed also in juveniles shells. Colours of protoconch, teleoconch and base very constant in all speci- mens observed. Etymology. The species is named after Errina aspera (Linnaeus, 1767) the Hydrozoan Stylas- teridae whose beds characterize the type locality in Strait of Messina, Sicily. Distribution. Currently only known from the type locality. DISCUSSION After the institution of the genus Jujubinus by Monterosato (1884), in recent years an increasing number of studies have greatly contributed to a bet- ter knowledge of this group of small trochids, with the description of new species and the rediscovery of some not yet well understood ones (Bogi & Cam- pani, 2005; Spanu, 2011; Mariottini et al., 2013; Smriglio et al., 2014; Smriglio et al., 2015). With this note we described/, errinae n. sp. (Figs. 1-18, 37), so increasing the number of the typical Juju- binus species [i.e. shell with prosocline lamellae between the spiral threads of variable strength, often beaded (Monterosato, 1884)] to be quoted for the Italian coast. The new taxon has been compared Figures 1-3. Jujubinus errinae n. sp., holotype (MZB60155), 4.9 mm (H) x 4.0 mm (W), from type locality (Strait of Messina), 100 m depth. Figures 4-5. Jujubinus errinae n. sp., paratype A (SG), 6.0 mm (H) x 4.7 mm (W) from type locality (Strait of Messina), 100 m depth. 62 Carlo Smriglio etalii Figures 7-15. Jujubinus errinae n. sp., holotype, SEM analyses, details of the shell. Figures 16-18. Jujubinus errinae n. sp., Strait of Messina, paratype R, 1.8 m (H) x 1.9 mm (W), CS-PM collection. Subadult specimen with basal cord sculptured by very pronounced tubercles. A new Jujubinus from the Mediterranean Sea 63 with three species showing a similar sculpture and occurring in the near Sicily Channel, J. catenatus, J. montagui and J. tumidulus (Curini & Palazzi, 1982). In particular, J. errinae n. sp. differs from/. catenatus (Figs. 19-36), the most closely related species which has an evident “pear-shaped” shell outline and shows a stronger sculptured orna- men- tation of the spiral cord interspaces, as well as a different background colour, being uniformly red- dish-greenish in J. catenatus , while the spiral cords of J. errinae n. sp. are white-spotted producing a typical shell pattern of irregular and interrupted axial stripes. The new species differs from J. montagui for its lower ratio H/W, the sculpture more tuberculate and densely ornamented with growth striae, producing a more jagged appearance of the shell surface, and the different shell chro- matic pattern. The shell colour of J. montagui is generally whitish or greyish with irregular brown axial stripes and basal cords with equally spaced and alternate brown-white dashes (Scaperrotta et al., 2010). Jujubinus errinae n. sp. differs from J. tumidulus being greater in size, having a much stronger sculpture, higher ratio H/W and a different shell colour, which in the latter species is generally uniformly creamy-whitish with brown spotted spiral cords (Scaperrotta et al., 2009). Noteworthy, the new taxon shows in the initial teleoconch whorls the basal cord strongly rippled, which becomes flat in the following whorls. This morpho- logical feature, very evident in juvenile shells (Figs. 16-18), regularly disappears during the shell devel- opment (Figs. 7-15). More generally, J. errinae n. sp. differs from most of the Atlantic and Mediter- Figures 19-21 .Jujubinus catenatus Ardovini, 2006. Sicily Channel. Figures 22-24. Jujubinus catenatus. Sicily Channel. 64 Carlo Smriglio etalii Figures 25-32. Jujubinus catenatus Ardovini, 2006. Specimen of figure 19. Sicily Channel, SEM analyses, details of the shell. Figure 33. Jujubinus catenatus. Strait of Messina, CS-PM collection. Subadult specimen. Figures 34-36. Jujubinus catenatus. Specimen of figure 22. Sicily Channel, SEM analyses, details of the shell. A new Jujubinus from the Mediterranean Sea 65 ranean Jujubinus species by its strongly tuberculate and jagged teleoconch sculpture, with evident lamellae in the interspaces and for its diagnostic coloration (see Description), never observed in any Recent Jujubinus distributed in Atlantic Ocean and Mediterranean Sea. The new species is known currently so far only from the type locality, in the Strait of Messina, sug- gesting to be another new endemism for this area (Fig. 37). The Strait of Messina is a complex and diversified environment having in the tidal-induced upwelling its main physical constraint. The up- welling, causing nutrient enrichment and temper- ature lowering of surface water both supports ex- ceptionally dense populations of suspension feeders (Mistri & Ceccherelli, 1995; De Domenico et al., 2009) and allows the settlement of Pliocene Atlantic remnants (Fredj & Giaccone, 1995). In this area, hard substrate corresponding to the Colantoni et al. (1981) “ rough bottoms with pinnacles' 1 ’, are charac- terized by dense and extensive colonies of the Hydrozoan Stylasteridae Errina aspera, known only for Gibraltar and the Messina Straits, which hosts an abundant and peculiar benthic fauna of Atlantic origin (Giacobbe & Spano, 2001; Giacobbe et al., 2007). Such well-known associated fauna was found in the sampled E. aspera beds (DG001 and DG04; 90-115 m depth), together with less frequent “accessory” species, as the bivalve Spondylus gussoni O.G. Costa, 1829, and the here described J. errinae n. sp. Such associated fauna was also found deeper (St 02; 201 m depth), on partially consolidated coarse sediment, colonized by E. aspera together with the giant barnacle 4 - Figure 37. Distribution of Jujubinus errinae n. sp. Pachylasma giganteum (Philippi, 1836). Differ- ently, the bathyal bottom sediment collected on the Gioia Basin slope (St IB; 371 m), characterized by terrigenous gravelly sands, showed a mixture of autochthonous (bathyal) and allochthonous (sub- tidal) bioclastic remains, which included J. errinae n. sp. specimens. Interestingly, in the same geo- graphical area is present Jujubinus curinii Bogi et Campani, 2005, another endemism belonging to the so-called “smooth” Jujubinus complex (Smriglio et al., 2014 and references therein). Such co-occur- rence of congeneric endemisms is not surprising, since the two species are living in different habitats whose peculiarities have been put in evidence in literature. Jujuubinus errinae n.sp., as accessory species in the E. aspera assemblages, might rep- resent a further Atlantic relict fauna having in the Strait of Messina its areal distribution. ACKNOWLEDGEMENTS Sincere thanks are due to Andrea Di Giulio and Patrizio Tratzi (Dipartimento di Scienze, Universita “Roma Tre”, Rome, Italy), for the SEM photo- graphs carried out at the LIME. We would like to express our gratitude to Massimo Appolloni (Museo Civico di Zoologia, Rome, Italy) for the ex- amination of the Jujubinus material kept in the Monterosato collection. A special thanks to Walter Renda (Reggio Calabria, Italy) for his great and disinterested help in this research. REFERENCES Barbero Bianchi C.N., 2004. Proposta di suddivisione dei mari italiani in settori biogeografici. Notiziario SIBM, 46: 57-59. Bogi C. & Campani E., 2005. Jujubinus curinii n. sp. una nuova specie di Trochidae per le coste della Sicilia. Bollettino Malacologico, 41: 99-101. 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Giacobbe S., Laria G. & Spano N., 2007. Hard bottom assemblages in the strait of Messina: distribution of Errina aspera L. (Hydrozoa: Stylasteridae). Rapports Commission International Mer Mediterranee, 38: 485. Gofas S. & Bouchet P., 2015. Jujubinus. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php7pMaxde- tails&id=138591 on 2015-09-12. Mariottini P., Di Giulio A., Apolloni M. & Smriglio C., 2013. Phenotypic diversity, taxonomic remarks and updated distribution of the Mediterranean Juju- binus baudoni (Monterosato,1891) (Gastropoda Trochidae). Biodiversity Journal, 4: 343-354. Mistri M. & Ceccherelli V.U., 1995. Produzione sec- ondaria di organsmi coloniali di substrato duro: i gorgonacei. S.IT.E. atti, 16: 463-465. Monterosato T.A. di Maria di, 1884. Conchiglie littorali mediterranee. Naturalista Siciliano, Palermo, 3: 102— 111 . Scaperrotta M., Bartolini S. & Bogi C., 2009. Accresci- menti. Stadi di accrescimento dei Molluschi marini del Mediterraneo. I. L'lnformatore Piceno, Ancona, 167 pp. Scaperrotta M., Bartolini S. & Bogi C., 2010. Accresci- menti. Stadi di accrescimento dei Molluschi marini del Mediterraneo. II. L'lnformatore Piceno, Ancona, 176 pp. Smriglio C., Di Giulio A. & Mariottini P., 2014. Descrip- tion of two new Jujubinus species (Gastropoda: Trochidae) from the Sicily Channel with notes on the Jujubinus curinii species complex. Zootaxa, 3815: 583-590. Smriglio C., Mariottini P. & Oliverio M., 2015. A new species of the Jujubinus curinii species complex: J. alboranensis n. sp. (Gastropoda: Trochidae) from the Alboran Sea. Iberus, 33: 151-157. Spanu M.T., 2011. Prima segnalazione di Jujubinus baudoni (Monterosato, 1 89 1 ex H. Martin ms) (Gast- ropoda: Trochidae) per la Sardegna e le acque italiane. Bollettino Malacologico, 47: 135-137. Biodiversity Journal, 2016, 7 (1): 67-78 Monograph Check-list of the Nudibranchs (Mollusca Gastropoda) from the biodiversity hot spot “Scoglio del Corallo” (Argentario promontory, Tuscany) Giulia Furfaro* & Paolo Mariottini Dipartimento di Scienze, Universita “Roma Tre”, Viale G. Marconi 446, 1-00146 Rome, Italy ’Corresponding author: giulia.furfaro@uniroma3.it ABSTRACT The Mediterranean nudibranch (Mollusca Gastropoda) fauna is part of complex communities belonging to the Mediterranean endemic “Coralligenous”. This important ecosystem shows a high species richness and functional diversity with assemblages of species tied together by major trophic and ecological relationships. A first check-list for the biodiversity hot spot “Scoglio del Corallo”, located along the coast of the Argentario promontory (Tuscany, Tyrrhenian Sea) is here reported. KEY WORDS Nudibranchs; biodiversity; check-list; Tyrrhenian Sea. Received 28.01.2016; accepted 27.02.2016; printed 30.03.2016 Proceedings of the Ninth Malacological Pontine Meeting, October 3rd-4th, 2015 - San Felice Circeo, Italy INTRODUCTION Nudibranchs are molluscs brightly coloured and frequently photographed by Scuba diver amateurs, since these sea slugs can be found in most coastal areas of the world, from polar to tropical waters. The most Nudibranchia diversity is known for shallow waters, ranging 0-30 m depth, but deep-sea research is unravelling high levels of previously unknown diversity of these molluscs at high depths too (Valdes, 2008; Oskars et al., 2015). The Mediter- ranean nudibranch diversity to date is of about 270 species, according to more recent checklists (Oztiirk et al., 2014; Trainito & Doneddu, 2014), regional faunal catalogues and Internet forum (WoRMS, Sea-slug forum). Although Mediterranean nud- ibranch species richness is smaller than that of the Indo-Pacific biogeographic region or the Caribbean Sea (Atlantic Ocean), the Mediterranean fauna has a high level of endemic diversity. Their scarce mobility in some cases leads them to live their entire life cycle associated to their trophic source and this is the reason why they are deeply related to the most important endemic habitats of the semi- closed Mediterranean Sea. In fact, they are common inhabitants of the Mediterranean benthic ecosystem defined as “Coralligenous” (Ballesteros, 2006), where they livefeeding on a broad range of different substrates (Sponges, Cnidarians, Bryozoans, Tunicates and other sessile animals) (Gutierrez, 2015). These complex communities are composed of a wide variety of suspension feeders, exhibiting high species richness and functional diversity (Gili & Coma, 1998). Recent molecular studies (e.g., Schrodl et al., 2011; Zapata et al., 2014) have proposed a new classification on the base of the polyphyly showed by this group that nowadays is split into 3 different Suborders (WoRMS: Gofas, 2015). This work aims to produce the first compre- hensive catalogue of the nudibrachs for the Biod- 68 Giulia Furfaro & Paolo Mariottini iversity hot spot “Scoglio del Corallo”, located along the coast of the Argentario Promontory (Tuscany, Tyrrhenian Sea) (Figs. 1-3), based on a fieldwork carried out by the authors in the last two years. An annotated Nudibranch checklist is pro- duced discussing taxonomic problems and new eco- logical data (association with other organisms, parasitism, cryptic species and geographical distri- bution) whenever relevant. Each species observed has been photographed in field and ecological and di- stribution data are provided for all species recorded. MATERIAL AND METHODS Sampling area “Scoglio del Corallo” is an underwater rocky hab- itat located in the in National Park of “Arcipelago Toscano” (42°23’60.00”N, 11°5 , 30.00”E, Central Tyrrhenian Sea). This submarine formation out- crops just for a few centimetres (depending on the marine tide) from the surface and slopes down vertically to a depth of 35 meters (Figs. 3,4). The most relevant inhabitant of this area is the Octocoral Corallium rubrum (Linnaeus, 1758) (Cnidarian), a Mediterranean endemic species included in several European and International protocols for conserva- tion, like the FAO General Fisheries Commission for the Mediterranean (GFCM) and the Convention on International Trade in Endangered Species (CITES). The presence of C. rubrum seems to be closely related to the high level of biodiversity characteristic of this area (Gili & Coma, 1998) (Figs. 5-9). This site is very small in extent (about 500 m 2 ), but nevertheless characterized by a set of rocks and walls forming canyons, caves and platforms placed on a muddy substrate creating a lot of microhabitats where a conspicuous number of species live and reproduce (Fig. 5). Protocol Sampling Sampling took place between the years 2013 to 2015 as a part of a broader research project (“Pro- ject Baseline Corallium rubrum ”, directed by the “Global Underwater Explorer” No-Profit Organiza- tion) aiming to produce the first characterization of this biota and of its associated biocoenoses. The produced preliminary data will become the starting Figures 1-3. Study area. Location of the “Scoglio del Corallo” (“Arcipelago Toscano”, 42°23’60.00”N, 1 1°5’30.00”E, Central Tyrrhenian Sea) in the Mediter- ranean Sea. Check-list of the Nudibranchs from the biodiversityhot spot “Scoglio del Corallo ” (Argentario promontory, Tuscany) 69 point for monitoring future environmental changes and to evaluate possible conservation strategies. Materials were sampled using SCUBA diving techniques. Specimens were obtained by manual collecting, photographed and fixed for future DNA extraction and anatomical studies in 96 % ethanol. Some species were observed and photographed on their natural habitat during field campaigns, but not captured. RESULTS AND DISCUSSION For the first time a Nudibranchs catalogue from a Tyrrhenian Sea submarine hot spot of biodiversity is here provided. A total of 23 species of nud- ibranchs belonging to 9 different families were col- lected during the project. Among these, 4 are endemics of the Mediterranean Sea showing the importance of this Mediterranean coralligenous as- semblage. All collected species coexist in this small area according to the high biodiversity showed by this hot spot marine site. The list of the sampled species is here reported, with notes on their ecology and distribution according to OPK-Opistobranquis (Available from http://opistobranquis.info/en/), Sea Slug Forum (Australian Museum, Sydney, Avail- able from http://www.seaslugforum.net/), World Register of Marine Species (WoRMS, Available from http://www.marinespecies.org at VLIZ), ‘'Sea slug of the Algarve” (Calado & Silva, 2012), “Nud- ihranchi del Mediterraneo ” (Trainito & Doneddu, 2014) and personal underwater observations. Phylum MOLLUSCA Classis GASTROPODA Subclass HETEROBRANCHIA Infraclass OPISTHOBRANCHIA Order NUDIBRANCHIA Suborder DEXIARCHIA Infraorder CLADOBRANCHIA Parvorder AEOLIDIDA Familia FACELINIDAE Bergh, 1889 Genus Craten a Bergh, 1864 1. Cratena peregrina (Gmelin, 1791) (Fig. 10) Ecology. This species commonly feeds on hy- droids of the genus Eudendrium Ehrenberg, 1834 on which it usually lays eggs. Cratena peregrina lives between a few meters from the surface till about 50 meters depth. Distribution. It has been found from Western to Eastern basin of the Mediterranean Sea, in the Portuguese and Andalusian Atlantic coasts and in the Canary Islands. It was also informally recorded from Senegal, South Africa, India and in Western Atlantic. Genus Facelina Alder et Hancock, 1855 2. Facelina annulicornis (Chamisso et Eysenhardt, 1821) (Fig. 11) Ecology. This species has a varied diet con- sisting on different genera of Hydrozoans: Eu- dendrium Ehrenberg, 1834, Obelia Peron et Lesueur, 1810, Pennaria Goldfuss, 1820 and Tubu- laria Linnaeus, 1758. Distribution. WoRMS (2015) recorded it from Mediterranean Sea and Atlantic Ocean (Ireland, United Kingdom, Azores, Portugal). The recent work of Oztiirk et al. (2014) expands its distribution range to the Turkish coasts of Aegean Sea. 3. Facelina rubrovittata (Costa A., 1866) (Fig. 12) Ecology. The few pictures of this rare nud- ibranch often show it staying on algae substratum. On the diet of F. rubrovittata little is known but it seems to feed on hydrozoans as well as most of the aeolids do. Distribution. It is distributed from the whole Mediterranean Sea till the Atlantic coasts of Spain. Familia FLABELLINIDAE Bergh, 1889 Genus Calmella Eliot, 1910 4. Calmella cavolini (Verany, 1846) (Figs. 13, 14) Ecology. This aeolid species usually feeds on Halecium pusillum Sars, 1856 and Eudendrium racemosum (Cavolini, 1785) but can be found on different substrates. This small nudibranch can be easily misidentied with the sibling species Pisei- notecus gaditanus Cervera, Garcia-Gomez et Garcia, 1987 from which it can be recognized by the absence, on its cerata, of the little white spots typical of P. gaditanus. Interestingly we could observe some individuals (Fig. 14) with very few 70 Giulia Furfaro & Paolo Mariottini white dots, whose identification needs possibly a DNA barcoding approach. Distribution. This endemic species originally was found only in the western coast of Mediter- ranean Sea but on the base of recent records its dis- tribution range now includes also the Turkish coasts and the Atlantic coast of the Iberian Peninsula. Genus Flabellina Gray, 1833 5 . Flabellina affinis (Gmelin, 1791) (Fig. 15) Ecology. Flabellina affinis is a very common species present all year long often feeding on colon- ies of Eudendrium spp. and belongs to the complex of the ‘pink Flabellinidae species’, see below F. ischitana and F. pedata. This species usually co- exists in the same arborescent hydrozoan colony with C. peregrina and can be parasitized by Cope- pods of the family Splanchnotrophidae, whose eggs often can be seen extruding from the notum of the host. Distribution. This is one of the most common European species ranging it from the eastern coast of Mediterranean Sea to the Atlantic basin of Spain and Portugal and in the Canarias islands. 6 Flabellina babai Schmekel, 1972 (Fig. 16) Ecology. This species shows a large body size atypical for a common ‘Flabellinid’. It can be found easily on different substrates mostly on hydroids of the genus Campanularia Lamarck, 1816, Eu- dendrium Ehrenberg, 1834 and Bougainvilla Lesson, 1 830, but is still unclear if it feeds on them. Distribution. This species has been recorded throughout the Mediterranean Sea and also in Senegal. 7. Flabellina ischitana Hirano et Thompson, 1990 (Fig. 17) Ecology. This species feeds on two different species of athecate hydrozoans of the genus Eu- dendrium , i.e. E. racemosum (Cavolini, 1785) and E. glomeratum Picard, 1952 often coexisting with F. affinis. They are morphologically very similar and easily confused with each other and, as mentioned above, both belong to the complex of the ‘pink Flabellinidae species’. Distribution. Its distribution overlap with the geographical range of F. affinis going from eastern basin of Mediterranean sea to the Atlantic coast of Iberian peninsula. 8. Flabellina lineata (Loven, 1846) (Fig. 18) Ecology. Mediterranean specimens of this ‘Fla- bellinid’ usually feed on Eudendrium spp. while the extra-Mediterranean individuals were observed on different species of hydroids like Tubularia indivisa Linnaeus, 1758, Coryne eximia Allman, 1859, Hy- drallmania falcata (Linnaeus, 1758) and Sertularia argen tea Linnaeus, 1758. Distribution. This species is distributed in the Atlantic Ocean, from the Arctic Circle to the French Atlantic coast, and in the European waters. 9. Flabellina pedata (Montagu, 1816) (Fig. 19) Ecology. Flabellina pedata also belongs to the complex of the ‘pink Flabellinidae species’, see above, being very similar to F. affinis and F. is- chitana from which differs on the base of possess- ing single cerata, not clustered together on a single peduncles, and by a smoothed rhinophores. It feeds on athecate hydrozoans of genus Eudendrium (especially in the Mediterranean Sea), but also on sertularids of genus Abietinaria Kirchenpauer, 1884 and on the plumularid genus Aglaophenia Lamour- oux, 1812. Recently was discover a new species of Flabellinid, Flabellina albomaculata Pola, Carmona, Calado et Cervera, 2014, very similar to F. pedata and easily confused with it. Distribution. This common Flabellinid is dis- tributed from eastern basin of the Mediterranean Sea to the Strait of Gibraltar and in the Atlantic Ocean from the Azores to the North Atlantic Coast of Norway. Parvorder CLADOBRANCHIA Familia PROCTONOTIDAE Gray, 1853 Genus Janolus Bergh, 1884 10. Janulus cristatus (Delle Chiaje, 1841) (Fig. 20) Ecology. This species lives between 10 and 40 m deep on a rocky substrate. Usually it was asso- ciated to different Bryozoans on which J. cristatus seemed to feed on. Alcyonidium gelatinosum (Hud- Check-list of the Nudibranchs from the biodiversityhot spot “Scoglio del Corallo ” (Argentario promontory, Tuscany) 7 1 son) J.V. Lamouroux, Bicellariella ciliata (Lin- naeus, 1758), Bugulina avicularia (Linnaeus, 1758), B. flabellata (Thompson in Gray, 1848), B. turbinata (Alder, 1857), Bugula neritina (Linnaeus, 1758) and species of genus Cellaria Ellis et Solan- der, 1786 were cited as a presumed preys. Distribution. It is commonly found throughout the Mediterranean Sea and in the North-eastern Atlantic Ocean from Norway to Morocco coasts. Parvorder DENDRONOTIDA Familia TRITONIIDAE Lamarck, 1809 Genus Marionia Vayssiere, 1877 11 .Marionia blainvillea (Risso, 1818) (Fig. 21) Ecology. It is recorded to feed on different preys like Alcyonium acaule Marion, 1878, A. pal- matum Pallas, 1766, Eunicella cavolinii (Koch, 1887), E. singularis (Esper, 1791), Eunicella sp., Leptogorgia sarmentosa (Esper, 1789), Para- muricea clavata (Risso, 1826). The juveniles have different body colours; in particular they are com- pletely white while the adults range in colour from a pale translucent orange to a deeper reddish brown with irregular white patches. They can be parasit- ized by Copepods like the ectoparasitic Doridicola comai Conradi, Megina et Lopez-Gonzalez, 2004 and the endoparasitic Linaresia bouligandi Stock, 1979 and L. mammillifera Zulueta, 1908. Distribution. Its geographical range goes from the whole Mediterranean Sea to the North-eastern and South-eastern Atlantic Ocean (Angola). Genus Tritonia Cuvier, 1798 12. Tritonia manicata Deshayes, 1853 (Fig. 22) Ecology. This species lives in shallow and very bright waters between the rhizomes of Posidonia oceanica (Linnaeus) Delile, 1 8 1 3 or on a rocky sub- strates where it can find a lot of Anthozoan (Cnid- aria) species. The Stoloniferous group is the one on which T. manicata seems to feed on, in particular on genus Cornularia Lamarck, 1816 and Clavu- laria Greville, 1865. Distribution. Present along the coasts of the Mediterranean Sea and also recorded from coast of Morocco and North- Atlantic British islands. 13. Tritonia striata Haefelfmger, 1963 (Fig. 23) Ecology. This species lives in shallow waters on rocky substrates full of algae, sponges and cnid- arians. It has been recorded to feed on the soft coral Paralcyoniums pinulosum Delle Chiaje, 1822. Distribution. Tritonia striata is known to be endemic of the Mediterranean Sea but recently it has been also recorded from the Gulf of Biscay in North Atlantic Ocean. Suborder EUCTENIDIACEA Infraorder DORIDACEA Familia ONCHIDORIDIDAE Gray, 1827 Genus Diaphorodoris Iredale et O'Donoghue, 1923 14. Diaphorodoris papillata Portmann et Sand- meier, 1960 (Fig. 24) Ecology. It feeds on Bryozoans so it is often observed in habitats rich in algae and sessile inver- tebrate fauna. Distribution. This species is endemic of the Mediterranean Sea but recorded also from coasts of Portugal and Strait of Gibraltar. 15. Diaphorodoris luteocincta var. alba (M. Sars, 1870) (Fig. 25) Ecology. It is reported to feed on different bryo- zoans genus Smittina Norman, 1903, Cellepora Linnaeus, 1767 and Crisia Lamouroux, 1812. It can be found in a rock walls hosting bryozoans, scyaphilic algae, hy droids and sponges. Distribution. There are two different morpho- types referring to D. luteocincta var. alba and D. luteocincta var. reticulata on the base of a dorsal notum completely white (var. alba) or red coloured (var. reticulata). These two morpho variants share the same wide distribution inhabiting the Mediter- ranean Sea and North-Eastern Atlantic Ocean (Trainito & Doneddu, 2014). Familia POLYCERIDAE Alder et Hancock, 1845 Genus Polycera Cuvier, 1816 16. Polycera quadrilineata (O. F. Muller, 1776) (Fig. 26) Ecology. Different species of Bryozoans were reported to be the substrate (possibly food) of the 72 Giulia Furfaro & Paolo Mariottini P. quadrilineata. This species lives in a rocky hab- itats where is relatively common. This species is often parasitized by Copepod Crustaceans belong- ing to the genus Splanchnotrophus Hancock et Norman, 1863 with the injection of its eggs into the body tissues of its host. Distribution. This species is distributed in Western Europe from Iceland and Greenland to the entire Mediterranean Sea. Familia CHROMODORIDIDAE Bergh, 1891 Genus Felimare Ev. Marcus et Er. Marcus, 1967 17. Felimare fontandraui (Pruvot-Fol, 1951) (Fig. 27) Ecology. Felimare fontandraui feeds on Sponges belonging to the genus Dysidea Johnston, 1842. This species can be found during all the year from the intertidal zone to about forty meters deep. It is very variable in colour morphs and some specimens can be misidentified with the sister species Feli- mare tricolor (Cantraine, 1835) from which can be recognized by the presents of a white basal spots on the rhinophores and other diagnostic characters. Distribution. Its distribution ranges from both the eastern and western Mediterranean basins to the North-eastern Atlantic coasts. 18. Felimare picta (Schultz in Philippi, 1836) (Fig. 28) Ecology. This species feeds on different sponges like species belonging to genus Ircinia Nardo, 1833, Crella Gray, 1867 and Dysidea Johnston, 1842. This common species lives on rocky substrate and shows different colour morphotypes described in the past like different subspecies. Distribution. Felimare picta has a wide spread distribution. It lives from the western coast of the Atlantic Ocean, Brazil and Florida, to the eastern Atlantic, Spanish and African coast as well and in the entire Mediterranean Sea. 19 . Felimare tricolor (Cantraine, 1835) (Fig. 29) Ecology. Felimare tricolor lives in rocky sub- strates from intertidal zone to about hundred meters deep. It feeds on different genera of sponges; Dysidea Johnston, 1842, Scalarispongia Cook et Bergquist, 2000 and Spongia Linnaeus, 1759. Distribution. This common species is distribu- ted in the Mediterranean Sea and in the North-ea- stern Atlantic Ocean. Genus Felimida Ev. Marcus, 1971 20 Felimida krohni (Verany, 1846) (Fig. 30) Ecology. This species has a morphology si- milar to the sister species Felimida britoi (Ortea& Perez, 1983), from which can be recognized by differences in the shape of the rhinophores and the mantle colour pattern. Its bathymetric range goes from subtidal zone down to 50 meters depth where it lives on different sponges like Hymenia- cidon perlevis (Montagu, 1814) and species of genus Ircinia Nardo, 1833. Its diet is still not clear. Distribution. The distribution of this species goes from the eastern basin of the Mediterranean Sea to the Strait of Gibraltar. It lives also in the North eastern Atlantic Ocean from Canary Islands and north coasts of Africa to the Atlantic coasts of Spain and France. 21. Felimida luteorosea (Rapp, 1827) (Fig. 31) Ecology. It lives under stones and on illumin- ated precoralligenous walls from 10 to 50 meters deep. It is reported to feeds on sponges like Aplysilla rosea (Barrois, 1876) and Spongionella pulchella (Sowerby, 1804). Distribution. It is distributed in the Mediter- ranean Sea and in the North Atlantic Ocean from the north coast of France and Spain to Angola and Canary islands. Familia DISCODORIDAE Bergh, 1891 Genus Peltodoris Bergh, 1880 22. Peltodoris atromaculata Bergh, 1880 (Fig. 32) Ecology. This common nudibranch lives on rocky bottoms usually associated to its prey, the sponge Petrosia ( Petrosia ) ficiformis (Poiret, 1789). It is extremely abundant in the coralligenous where it lives searching for its food or staying on it. This sea slug is very sedentary so it can be found on the same sponge for different days. Check-list of the Nudibranchs from the biodiversityhot spot “Scoglio del Corallo ” (Argentario promontory, Tuscany) Figures 4-9. Underwater photographs of the “Scoglio del Corallo”, showing the Cor allium rubrum assemblages. 74 Giulia Furfaro & Paolo Mariottini Figures 10-15. Fig. 10: Cratena peregrina (Gmelin, 1791). Fig. 11: Facelina annulicornis (Chamisso et Eysenhardt, 1821). Fig. 12: Facelina rubrovittata (Costa A., 1866). Figs. 13,14: Calmella cavolini (Verany, 1846). Fig. 15: Flabellina affinis (Gmelin, 1791). Check-list of the Nudibranchs from the biodiversityhot spot “Scoglio del Corallo” (Argentario promontory, Tuscany) 75 Figures 16-21. Fig. 16: Flabellina babai Schmekel, 1972. Fig. 17. Flabellina ischitana Hirano et Thompson, 1990. Fig. 18. Flabellina lineata (Loven, 1846). Fig. 19: Flabellina pedata (Montagu, 1816). Fig. 20: Janulus cristatus (Delle Chiaje, 1841). Fig. 21: Marionia blainvillea (Risso, 1818). 76 Giulia Furfaro & Paolo Mariottini Figures 22-27. Fig. 22: Tritonia manicata Deshayes, 1853. Fig. 23: Tritonia striata Haefelfmger, 1963. Fig. 24: Diaphorodoris papillata Portmann et Sandmeier, 1960. Fig. 25: Diaphorodoris luteocincta var. alba (M. Sars, 1870). Fig. 26: Polycera quadrilineata (O. F. Muller, 1776). Fig. 27: Felimare fontandraui (Pruvot-Fol, 1951). Check-list of the Nudibranchs from the biodiversityhot spot “Scoglio del Corallo” (Argentario promontory, Tuscany) 77 Figures 28-33. Fig. 28: Felimare picta (Schultz in Philippi, 1836). Fig. 29. Felimare tricolor (Cantraine, 1835). Fig. 30: Felimida krohni (Verany, 1846). Fig. 31: Felimida luteorosea (Rapp, 1827). Fig. 32: Peltodoris at- romaculata Bergh, 1880. Fig. 33: Phyllidia flava Aradas, 1847. 78 Giulia Furfaro & Paolo Mariottini Distribution. This is one of the most common species of the Mediterranean Sea. It is also recorded from Western Atlantic Ocean from Portuguese coasts to Canary Islands. Familia PHYLLIDIIDAE Rafinesque, 1814 Genus Phyllidia Cuvier, 1797 23 . Phyllidia flava Aradas, 1847 (Fig. 33) Ecology. This interesting sea slug has a charac- teristic body colour that can camouflage it when it is associated to sponges like Axinella cannabina (Esper, 1794), A. polypoides Schmidt, 1862 and Acanthella acuta Schmidt, 1862. It has been known to feed on the latter sponge. Distribution. This species is rare and distrib- uted throughout the Mediterranean Sea, it has been also recorded from the Canary Islands. AKNOWLEDGEMENT The authors gratefully acknowledge the GUE instructor Luca Malentacchi (Arezzo, Italy), the “Project Baseline Cor allium rubrum ” manager, for its help during sampling and monitoring and for providing underwater photographs and videos. Thanks to the big community of volunteers that work for “Project Baseline Corallium rubrum ”. We are indebted to Monica Valdambrini (Arezzo, Italy), Massimiliano Falleri, Massimiliano Orsi and Gianluca Cireddu (Rome, Italy, Italy) for providing underwater photographs. We thank people from “Gruppo Malacologico Mediterraneo” (Rome, Italy) for their assistance during samplings. GF and PM wish to thank the University of Roma Tre for financial funding. REFERENCES Ballesteros E., 2006. Mediterranean coralligenous as- semblages: a synthesis of present knowledge. Ocean- ography and Marine Biology: An Annual Review, 44: 123-195. 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A new phylogeny of the Cephalaspidea (Gastropoda: Heterobranchia) based on expanded taxon sampling and gene markers. Molecular Phylogenetics and Evolution, 89: 130-150, doi.org/10.1016/j.ympev. 2015.04.011. OztiirkB., Dogan A., Bitlis-Bakir B. & Salman A., 2014. Marine molluscs of the Turkish coasts: an updated checklist. Turkish Journal of Zoology, 38: 832-879. doi:10.3906/zoo-1405-78 Schrodl M., Jorger K., Klussmann-Kolb A. & Wilson N.G., 2011. Bye bye "Opisthobranchia"! A review on the contribution of mesopsammic sea slugs to euthyneuran systematics. Thalassas, 27: 101-112. Sea Slug Forum, Australian Museum, Sydney, Available from http ://www. seaslugforum.net/ Trainito E. & Doneddu M., 2014. Nudibranchi del Mediterraneo. Ed. 11 Castello, Milano, 192 pp. Valdes A., 2008. Deep sea “cephalaspidean” hetero- branchs (Gastropoda) from the tropical southwest Pacific. In: Heros V., Cowie R.H. & Bouchet P. (Eds.), Tropical Deep-Sea Benthos 25. Memoires du Museum national d’Histoire naturelle, 196: 587-792. WoRMS Editorial Board, 2015. World Register of Marine Species. Available from http://www.marine- species. org at VLIZ. Accessed 2015-11-17. Zapata F., Wilson N.G., Howison M., Andrade S.C.S., Jorger K.M., Schrodl M., Goetz F.E., Giribet G. & Dunn C.W., 2014. Phylogenomic analyses of deep gastropod relationships reject Orthogastropoda. Proceedings of the Royal Society B: Biological Sciences, 281 (1794): 1-9. http://dx.doi.org/10.1098/ rspb. 2014.1739 Biodiversity Journal, 2016, 7 (1): 79-88 Monograph Barycypmea teulerei (Cazenavette, 1 845) (Gastropoda Cypraeidae): a successful species or an evolutionary dead-end? Marco Passamonti Dipartimento di Scienze Biologiche Geologiche e Ambientali (BiGeA), Via Selmi 3, 40126 Bologna, Italy; email: marco. passamonti@unibo.it ABSTRACT Barycypraea teulerei (Cazenavette, 1845) (Gastropoda Cypraeidae) is an unusual cowrie species, showing remarkable adaptations to an uncommon environment. It lives intertidally on flat sand/mud salt marshes, in a limited range, in Oman. On Masirah Island, humans probably drove it to extinction because of shell collecting. A new population, with a limited range, has recently been discovered, and this article describes observations I made on site in 2014. Evolution shaped this species into a rather specialized and successful life, but has also put it at risk. Barycypraea teulerei is well adapted to survive in its habitat, but at the same time is easily visible and accessible to humans, and this puts it at high risk of extinction. Evolution is indeed a blind watchmaker that ‘ has no vision, no foresight, no sight at all' . And B. teulerei was just plain unlucky to encounter our species on its journey on our planet. KEY WORDS Cypraeidae; Barycypraea teulerei ; Biology; Evolution; Blind Watchmaker. Received 23.02.2016; accepted 22.03.2016; printed 30.03.2016 Proceedings of the Ninth Malacological Pontine Meeting, October 3rd-4th, 2015 - San Felice Circeo, Italy INTRODUCTION “ Natural selection, the blind, unconscious, auto- matic process which Darwin discovered, and which we now know is the explanation for the existence and apparently purposeful form of all life, has no purpose in mind. It has no mind and no mind's eye. It does not plan for the future. It has no vision, no foresight, no sight at all. If it can be said to play the role of watchmaker in nature, it is the blind watch- maker". R. Dawkins, The Blind Watchmaker, 1986. Barycypraea teiderei (Cazenavette, 1845) (Gast- ropoda Cypraeidae) (Fig. 1) is one of only two relic species of the genus Barycypraea Schilder, 1927, along with the South African Barycypraea fultoni (Sowerby III, 1899) (Fig. 2). This genus is charac- terized by squat, heavy shells with a roughly trian- gular/pyriform shape. The mantle is always thin and almost transparent, whitish or pale brown, with little (. B . fultoni ) or no papillae (B. teulerei ). Both species appear to be well adapted to sand and/ or mud bottoms, although at very different depths. Barycypraea fultoni is a deep water species (Ber- gonzoni, 2012) while, as we will see in detail, B. teiderei is an intertidal one. The genus comprises few fossil species, among them B. ziestmani Liltved et Le Roux, 1988 from the Alexandria Formation (Neogene), Port Eliza- beth, S. Africa (Liltved, 2000), and the B. caputvi- perae species-complex from Indonesia (Miocene). The genus Barycypraea is morphologically and ge- netically linked to the genus Zoila Jousseaume, 80 Marco Passamonti 1884, which is endemic to Western and Southern Australia. In this sense, the entire evolution of this cowrie lineage has always been strictly related to the Indian Ocean basin. The supposed similarity to the Venezuelan/Colombian Muracypraea mus (Linnaeus, 1758) and other allied fossil species of the Caribbean genus Siphocypraea Heilprin, 1897 [f.i. S. problematica (Heilprin, 1887)], seems not fully supported by molecular studies (Meyer, 2004). A STORY OF A ONCE RARE SPECIES Barycypraea teulerei was once an extremely rare species. In 1964, only 35 specimens were present in European collections (Schilder, 1964). Since 1969, only guesses were available about its distribution and habitat, since no one had ever reported a precise locality for the species. Speci- mens were labeled from different localities, includ- ing the Persian Gulf, Hormuz Strait, Aden, Arabian Sea, Red Sea, Port Sudan etc. (see Scali, 2013 for a detailed list), but in fact, no one knew where this species came from. In March 1969, the very first specimens of B. teulerei were collected at Masirah Island (wrongly reported as Museera Island; Cross, 1969), a very re- mote island along the Oman coastline. Barycypraea teulerei appeared to live in very shallow water on sand/mud beds, and even outside the water during low tide. Since then, several malacologists made their way to Masirah to obtain specimens (see f.i. Williams, 1969; Luther, 1972; Charter, 1983) which soon became available for study and collection. One of the main sources was actually Dr. Donald ‘Don’ T. Bosch, who had a long service as a surgeon for the Sultanate of Oman. Dr. Bosch was the only surgeon in the entire country of 1 .5 million people, and contributed to the modernization of health care in Oman. In recognition of his achievements, the Sultan of Oman awarded him with the "Order of Oman" in 1972. Don was also an extensive shell collector and a pioneer of Oman malacology. Many Oman species have been named by or after him (e.g. Conus boschi Clover, 1972, Cymatium boschi Abbott et Lewis, 1970, etc.), and he also dedicated some to his wife Eloise (e.g. Acteon eloisae Abbott, 1970). Because of Don Bosch and other shell col- lectors, thousand of specimens were easily available for a while, and the species became quite common in collections. Eventually, in the early 90s, new fresh-collected B. teulerei began to disappear. By then, many col- lectors traveled to Masirah to collect specimens, without success. The species seemed to have simply vanished, probably due to over-collection and the relative ease of finding specimens by simply walking the flat beaches of the Island. Rumors were growing that this species had to be considered extinct. In December 2012, after some unsuccessful trips to Oman, B. teulerei was found again by Massimo Scali and his family along the coastline of Oman (Scali, 2013; 2014), in a locality kept secret since then. The population was very healthy, with several thousand specimens freely grazing on a muddy flat bottom. Again, it was confirmed that this species lives in the intertidal zone. At low tide, B. teulerei does not hide itself under stones, as most cowries would do, and it is quite often completely exposed. In figure 3 you can see some in situ specimens during the sygyzian tide of 2014. In December 2014, I was fortunate enough to join Massimo in his field trip to see this species on site. This article is basically a series of observations Eve made that I hope will be of interest to the readers. I will discuss some of the aspects of the biology of this species, and I will express some con- siderations about its evolution. THE HABITAT Once I arrived at the place during the sygyzial tide of December 2014, 1 soon realized we were in an unusual habitat for a cowrie species. What I saw was basically a muddy salt marsh, covered with patches of algal mats and a few dark gray and orange sponges (Figs. 4, 5). No rock or evident coral to be found for kilometers. The only available hard substrate was a few Pinna sp. standing out of the bottom and a few dead bivalve shells. The mud was very anoxic, dark colored and stinking of sul- fur. It was hard to walk on, at every step I remained glued in the mud. Despite this environment seeming quite inhospitable, a few minutes walking from the beach I found the first living B. teulerei. This mud flat is a relatively large area and we were walking, on average, 10-15 kilometers per day to observe B. Barycypraea teulerei (Gastropoda Cypraeidae): a successful species or an evolutionary dead-end? 81 Figure 1 . Barycypraea teulerei. Examples of variability of pattern of the species. Oman. Photo courtesy Massimo Scali and Beautifulcowries Magazine. Figure 2. Barycypraea fultoni. Examples of pattern and variability of the species. Mozambique and South Africa. Photo courtesy Mirco Bergonzoni and Beautifulcowries Magazine. 82 Marco Passamonti teulerei in situ. The other animals I was able to see were crabs, cuttlefishes, many bivalves, muricids, and some other cowrie species. Nevertheless, B. teulerei is by far the most common species in this environment. Its distribution is not even, however. Barycypraea teulerei tends to aggregate, and you can find dozens of specimens together in the same patch, then walk for minutes and not find one. What I observed is that the animals are active during the daytime, especially the small ones that I think may be males (see below). In many cases, they are heedless of being completely outside of the water. Walking on the flat for hours, I was also able to ob- serve a few species of sea birds including small waders, flamingos and seagulls. In comparing this to the previous known habitat of B. teulerei (which I indeed visited), the main dif- ference is that at Masirah Island the sediment is sand, and the bottom is not anoxic. In Masirah, the above-mentioned algal mats and sponges are nowa- days very rare, and the area looks more like a big sandy beach with scattered rocky patches. Never- theless historical records, as well as a few very dead shells, witness that the area once hosted B. teulerei. My guess is that, besides collection pressure, there could have been some environmental change. Figure 3. Barycypraea teulerei wandering on a mud flat outside water at sygyzial low tide in December 2014. Oman. Figure 4. The typical environment, at low tide, where Barycypraea teulerei are commonly found. Please note the algal patches. Oman. Figure 5. A close look of the sponge, common in the area, on which Barycypraea teulerei was seen eating. Oman. Barycypraea teulerei (Gastropoda Cypraeidae): a successful species or an evolutionary dead-end? 83 In my opinion, B. teulerei needs the presence of sponges to establish a healthy population. Dur- ing my observations, I was able to see a B. teulerei feeding on a dark gray/black sponge (Fig. 5), so I can confirm this species is spongivorous. How- ever, I cannot exclude it feeding on algae too, but I have not seen any doing so. This is another char- acteristic that joins B. teulerei to the spongivorous Zoila. REPRODUCTION AND LIFE CYCLE Two other things are, in my opinion, necessary for B. teulerei to establish a healthy population. Firstly, dead bivalve shells. Barycypraea teulerei uses these shells to nest its eggs. When a female is brooding eggs (as all cowries do), she hides herself and the eggs on the underside of the valve. Females are hidden by the bivalve shell, but you can spot them because several males are commonly found close to them (Fig. 6). Females, on average, tend to be bigger than males, although this is not always true. Massimo Scali also spotted a male fecundating a female on eggs (Fig. 7). This may be an indication that eggs are fecundated while females lay them, and a reason why males compete for laying females. Egg clusters are comprised of transparent cap- sules with brownish eggs inside. Immature capsules contain many eggs, but as development continues, only a few embryos per capsule are found. Embryos are easy to spot because they already have a formed shell (Fig. 8). Likely most of the eggs inside the capsule are only for embryo nutrition (nurse cells or intracapsular cannibalism?). It is therefore evident that this species has direct development and only one (or a few) newborns are hatching from each capsule. Direct development is, in cowries, considered an adaptation when a species depends on a limited food source (in this case sponges), so that newborns hatch close to their food source instead of being spread throughout wide areas as veligers. This direct development is again another similarity to Zoila. When we arrived in December, many speci- mens were brooding eggs and we seemed to be right in the middle of the reproductive season. Air temperatures in Oman during December are not ex- tremely hot, and during the day can reach 25-30°C. However, at night it can be as cold as 10°C or less. Figure 6. The typical behavior of a female breeding eggs. Above: the female is hidden under a dead bivalve shells, and two males are trying to fecundate. Below: same an- imals, after I turned the bivalve to make the female visible. Oman. Figur 7. A male Barycypraea teulerei fertilizes with his penis (A) a female that sits on a bivalve shell (B). Oman. Photo courtesy Massimo Scali. 84 Marco Passamonti Figure 8. Two views of egg clusters inside bivalve shells. Please note that each capsule may contain different numbers of brownish eggs, and as soon as the embryos get bigger, the number of them decreases. Intracapsular cannibalism? Oman. Photo courtesy Massimo Scali and Beautifulcowries Magazine. Our time at the site was basically the coldest part of the year, and I guess this is the main reason why B. teulerei reproduce during winter. This species is intertidal, so it is strongly influenced by solar heat and desiccation, and winter is the time of the year in which that is least likely to happen. The mud itself may also help in maintaining mollusk wetness and lowering temperature during air exposure. Moreover, water patches and little canals are still present in the mud flat, and some specimens (espe- cially males) seem to take refuge in these when the tide is very low. Finally, almost no specimen showed an expanded mantle, and this is certainly a behavior for retaining moisture and reducing de- hydration. Another surprising observation, confirmed by previous reports at Masirah, is that we couldn’t find any juvenile B. teulerei. All specimens were adults or, slightly sub-adult. Another important observa- tion is that while adults are very visible and active, sub-adults are more mimetic and tend to hide below the algal mats. The fact that no young B. teulerei were found points sharply to the possibility that this species has a synchronized life cycle, and all reproducing mollusks found are the ones born from eggs of the previous year. Moreover, another observation is important: although B. teulerei shells are very heavy, no shell seems gerontic and most of them are undamaged. It seems likely they had no time to be damaged, and maybe this is because all those reproducing shells are just one year old and reached sexual maturity only a few weeks before we arrived. If my hypothesis is correct, this would mean that B. teulerei is a cowrie with a very fast life cycle. Soon after December/January they hatch as small crawling snails. The snails, having thin shells, protect themselves from predators and desiccation by hiding inside the algal mats, which are actually quite intricate, and I guess these may also help in cooling the mollusks during the hot season low tides. Sponges are too small to be a suitable refuge even for the youngest snails. They develop this way until the beginning of the next reproductive season, when they complete development and start wander- ing for dead bivalve shells (if female) or other females (if males). Again, this peculiar life cycle, if confirmed, coincides quite remarkably with Zoila. Actually, Zoila newborns are very cryptic, as they hide inside sponges as protection from predation, and they only venture out into the open during reproductive season, when they reach adulthood and shells get thicker. Zoila friendii, for instance, broods eggs in the open (personal observation) just as B. teulerei does. The complete development of B. teulerei is there- fore spanning along the hot season. Oman is very hot during summer, easily reaching 40° C or more. I may imagine that, especially during low tide, the water could reach a very high temperature. Please Barycypraea teulerei (Gastropoda Cypraeidae): a successful species or an evolutionary dead-end? 85 Figure 9. Examples of the variability of the dorsum in Barycypraea teulerei, some resembling a false aperture. Oman. remember that this is a large lagoon flat and that the open ocean, which may be cooler, is quite distant. It is quite unbelievable, but apparently young B. teulerei are able to deal with these harsh conditions and reach adulthood with good success. A possib- ility is that B. teulerei migrates into shallow waters only to reproduce, and lives most of the year in deeper waters, where conditions are more stable. I do not think this is the case, because this species is not capable of fast movement, and the mud flat is several kilometers wide. Plus, I found most indi- viduals very far from the deeper areas (actually the closer we searched to the open ocean, the less spe- cimens were found). Moreover, it is unlikely that young B. teulerei are able to migrate back to the deeper water during their development, when they 86 Marco Passamonti are most vulnerable due to their thin shells. We also dredged for a few hours along the edge of the flat at a depth of 5-10 meters, and we found no B. teulerei , not even dead ones. Finally, what is the fate of the specimens that reached the first reproductive season? Do they survive to the next year? Is this species annual or not? Hard to say, but the fact that I could see very few damaged and no gerontic shells suggests that this species is rather annual, and after reproduction B. teulerei dies. If this is true, the population renews itself every year. It may seem strange that a mollusk forms such a hard, heavy shell in only one year, but there is no biological reason to disregard this hypothesis. On the other hand, it is true that we found very few dead shells, and there should be many more if they all die each year. It is also true that they can be easily burrowed into the soft mud bottom, so they would easily disap- pear. Nevertheless some dead shells are found beached as well. PREDATION How does B. teulerei deal with predators? As we mentioned, this species lives in open sand/mud flats, and they do not hide when adult. Moreover, several hundred specimens are found in relatively small areas. Barycypraea teulerei actually seems quite a successful species and, in fact, it is by far the most common cowrie in that particular habitat. Its behavior is quite the oppos- ite of other cowries inhabiting the same area, f.i. Naria turdus (Lamarck, 1810) and Palmadusta lentiginosa (J.E. Gray, 1825), which are found hiding inside the algal coating of the numerous Pinna sp. found on the muddy bottom. And this is not because they are smaller, since some local N. turdus may be as big as B. teiderei, and with a similarly thick shell. Among candidate predators, I may mention seagulls and crabs, which are common in the area, as well as other mollusks. However, very few shells (almost none) show signs of predation, and I have not seen any cracked shells on site or beached. Dead shells are also very rare, and when found, they do not show any sign of attack. It seems that predators are completely ignoring B. teulerei , an observation that was quite puzzling. Why should this species not be predated, and why does it actually seem to ignore predators? Is B. teulerei toxic, poisonous, or have a disgusting taste? Hard to say, but as far as I know, no toxic cowrie has ever been reported in literature. It is not unconceivable that perhaps they become toxic, repellent or disgusting by absorbing sub- stances from their food sponges. Only targeted chemical analyses would possibly solve this issue. Some clues may also come from the shell. As mentioned, B. teulerei has a very heavy shell. Its thickness is certainly an adaptation to prevent cracking by fishes, crabs or sea birds, as well as drilling by muricids or naticids. Moreover, its squat shape might also be an adaptation to per- fectly adhere it to the bottom (as in many other cowries). However, I may also argue that the peculiar pattern of the dorsum could have an ad- aptive function, although this is just a guess. In fact, even if the dorsum is characterized by a variable pattern (basically no two specimens are alike), most shells show a neat double dorsal line, framing a central groove, especially when the shell is thicker. More uncommonly, they show a dark blotch in the middle of the dorsum. Other kinds of patterns are rarer. In figure 9 you can see an over- view of the variability of the dorsal patterns. Con- trarily to the dorsum, the mouth is quite wide and uncolored. Considering all this, my guess is that the flashy dorsal color in this species might be either an aposematic coloration (in case the mol- lusk is toxic or has a bad taste), or, maybe, could represent a sort of ‘false aperture’ that may distract sea birds from the vulnerable parts of the animal. I may imagine seabirds being fooled and peck at the dorsum of the cowrie, which is actually a very hard part of the shell, completely disregarding the real aperture, where the mollusk would be more vulnerable. Of course this is just a guess, but it is of course not the first such case known in nature: for instance, you may find something similar in false eyes of fishes, which are adaptations to drive predators’ attacks to parts of the body that are less sensitive or critical for survival. Finally, please also note that this species has no teeth along its aperture, a characteristic that is very rare in cowries, even if some specimens may have some little denticles. Teeth in cowries have a par- ticular function, i.e. to narrow the aperture to pre- vent attacks from predators, since cowries have no operculum. Evidently this species has no need Barycypraea teulerei (Gastropoda Cypraeidae): a successful species or an evolutionary dead-end? 87 for teeth, and teeth, which are found in all other Barycypraea , are on their way to being lost. This is a very well known evolutionary process: no select- ive constraints (i.e. no need for teeth) allow accu- mulation of mutations, which result in the gene products having less or no function (i.e., the genes or the proteins involved in teeth production being partially or wholly inactivated). AN EVOLUTIONARY DEAD END? All this considered, B. teulerei shows a pleth- ora of remarkable adaptations to a very specific environment, which makes this species an outlier among cowries: i) it lives in the intertidal zone on sand/mud flats, where other cowries are rarer; ii) it is active during the day, at variance to other cowries; and iii) it is strictly dependent on a specific habitat and food source. Nevertheless it performs quite well when all these conditions are present, so we can say that this species appears very well adapted to its environment. Evolution has done “a good job” with this species. And, in fact, B. teulerei has no significant predators, at least when they are adult and freely grazing and mating in the open. On the other hand, its distribution range seems quite limited, maybe because of its specialist way of life. The absence of free-swimming larvae is certainly another concurring factor. We tried to find B. teulerei elsewhere along the Oman coast, with no success, although more research needs to be done. Unfortunately, the very limited distribution makes this species a highly vulnerable one. Actually the main concern for the survival of B. teulerei does not come from predators, but from humans. It was quite bad luck for B. teulerei to find a species collecting it in large numbers for its beauty, rather than for its taste. And it was bad luck indeed that this species is commercially valuable to collectors. The limited range does the rest. The story of the Masirah population teaches us that B. teulerei is indeed in high danger of extinction. That is why the new locality should absolutely remain secret, and I am not giving any precise indication as to where it is. It is also my opinion that this species should be protected by law. The life history of B. teulerei is, no doubt, a remarkable one. Evolution shaped this species to a rather specialized and successful life. At the same time, it has put B. teulerei at risk. Evolution is a blind process and of course it could not foresee that, at a certain point, this species would have en- countered another one: humans, predating it for its shine, beauty and striking colors. Evolution shaped B. teulerei to survive in its habitat, but at the same time made it so easily accessible to humans, and its highly specialized life puts it at risk of extinction. Evolution is indeed a blind watchmaker that ‘ has no vision, no foresight, no sight at alV (Dawkins, 1986). And B. teulerei was just plain unlucky to encounter our species during its journey on our planet. ACKNOWLEDGMENTS I wish to thank you very much Massimo Scali (Imola, Italy) for giving me the opportunity to visit and observe on site the new population of B. teulerei in Oman, and for providing some of the pictures. I also wish to thank very much the organ- izers of the VIII Pontine Malacological Congress, Silvia Alfinito and Bruno Fumanti (Sabaudia, Italy), as well as the Malalcos 2002 association and mem- bers, for inviting me to talk about B. teulerei. Finally, my acknowledgement goes to Federico Plazzi (Bologna, Italy) and Rex Stilwill (Grand Ra- pids, MI, USA), for their review of the manuscript. REFERENCES Bergonzoni M., 2012. Barycypraea fultoni, a tale of a fallen star. Beautifulcowries Magazine, 2:4-19. Charter B., 1983. Masirah, the teulerei island, revisited. Hawaiian Shell News, 31: 1,9. Cross E.R., 1969. The case history of a rare shell. Hawaiian Shell News, 17: 1,4. Dawkins R., 1986. The blind watchmaker. Norton & Company Inc., New York, U.S.A. Liltved W.R., 2000. Cowries and their relatives of southern Africa. II Edition revisited. Seacomber Publications, Cape Town, S. Africa, 232 pp. Luther F., 1972. A1 Masira - teuleri island. Hawaiian Shell News, 20: 1, 3. Meyer C.P., 2004. Towards comprensiveness: increased molecular sampling with Cypraeidae and its phylo- genetic implications. Malacologia, 46: 127-156. Schilder F.A., 1964. The true habitat of a rare cowry. Hawaiian Shell News, 12: 2. 88 Marco Passamonti Scali M., 2013. Barycypraea teulerei (Cazenavette, 1846). The rediscovery. Beautifulcowries Magazine 3: 4-11. Scali M., 2014. Barycypraea teulerei, going back to the recently discovered new population. Beautiful- cowries Magazine 5: 31-35. Williams M., 1969. Cypraea teulerei follow up. Letter from Arabia. Hawaiian Shell News, 17: 1,9. Biodiversity Journal, 2016, 7 (1): 89-92 Monograph Contribution to the knowledge of the molluscan thanato- coenosis of Zannone Island (Pontine Archipelago, Latium, ltaly).Additional reports Bruno Fumanti 1 & Italo Nofroni 2 'Via del Villaggio 108, 04010 Sabaudia, Latina, Italy; e-mail: bmno.fumanti@libero.it 2 Via Benedetto Croce 97, 00142 Rome, Italy; e-mail: italo.nofroni@uniromal.it "■Corresponding author ABSTRACT In this second paper concerning the molluscan fauna of Zannone Island (Pontine Archipelago, Italy) one sediment sample collected by scuba diving at a depth of 36.5 meters at SW of the isle was investigated. Altogether, 47 taxa, not yet reported for Zannone, were identified, brin- ging the total number of the molluscan thanatocoenosis of the island at 327 taxa. KEY WORDS Mollusca; thanatocoenosis; Zannone Island; Italy. Received 02.03.2016; accepted 18.03.2016; printed 30.03.2016 Proceedings of the Ninth Malacological Pontine Meeting, October 3rd-4th, 2015 - San Felice Circeo, Italy INTRODUCTION In a previous paper on the Mollusca of Zannone Island, Fumanti (2014) reported 280 malacological taxa. Recently prof. Riccardo Lubrano has provided one of us (I. N.) with a sediment sample collected on August 1 1th 2013 by means of scuba diving by Mr. Nino Baglio at 36.5 m depth SW of Zannone Island. The study of the sample has led to the iden- tification of several taxa, 47 of these, representing an increase of 14.4 % of the total, were not previ- ously reported. Finally, thanks to the finding of Spinoaglaja wil- pretii (Ortea, Bacallado et Noro, 2003) by Romani & Pagli (2014), the molluscan thanatocoenosis of the Isle of Zannone consists now of 327 taxa. Fur- thermore, in this paper, other species, indicated with an asterisk, already reported in Fumanti (2014) have been added with revised and updated nomen- clature recording the latest publication. RESULTS Taxonomic list Classis GASTROPODA Cuvier, 1797 Ordo PATELLOGASTROPODALindenberg, 1986 Familia LOTTED AE Gray, 1840 Genus Tectura Gray, 1 847 Tectura virginea (O.F. Muller, 1776) Ordo VETIGASTROPODA Salvini-Plawen, 1980 Familia SCISSURELLIDAE Gray, 1847 Genus Sinezona Finlay, 1926 Sinezona cingulata (O.G.Costa, 1861) Familia SKENEIDAE Clark, 1851 Genus Skenea Fleming, 1 825 Skenea serpuloides (Montagu, 1808) 90 Bruno Fumanti & Italo Nofroni Ordo CAENOGASTROPODA Cox, 1960 Familia SILIQUARIIDAE Anton, 1838 Genus Petalopoma Schiapparelli, 2002 Petalopoma elisabettae Schiapparelli, 2002 Familia SKENEOPSIDAE Iridale, 1815 Genus Skeneopsis Iridale, 1915 Skeneopsis planorbis (O. Fabricius, 1780) Familia JANTHINIDAE Lamarck, 1822 Genus Janthina Roding, 1798 Janthina pallida W. Thompson, 1840 Familia RISSOIDAE J.E. Gray, 1847 Genus Alvania Leach in Risso, 1826 Alvania dictyophora (Philippi, 1844) group Notes. Actually this species is under investiga- tion by Bruno Amati (Rome). Genus Setia H. Adams et A. Adams, 1852 Setia turriculata Monterosato, 1884 Familia CYPRAEIDAE Rafmesque, 1815 Genus Nana Broderip, 1837 Naria spurca (Linnaeus, 1758) Familia LIMACINIDAE Gray, 1840 Genus Thielea Strebel, 1908 Thielea inflata (d’Orbigny, 1836) Genus Limacina Bose, 1817 Limacina trochifovmis (d’Orbigny, 1836) Familia PERACLIDAE Tesch, 1913 Genus Peracle Forbes, 1844 Peracle reticulata (d’Orbigny, 1836) Familia MURICIDAE Rafmesque, 1815 Genus Dermomurex Monterosato, 1890 Dermomure x scalaroides (Blainville, 1826) Genus Murexsul Iridale, 1915 Murexsul aradasii (Monterosato in Poirer, 1883) Familia MITROMORPHIDAE Casey, 1904 Genus Mitromorpha Carpenter, 1865 *Mitromorpha columbellaris (Scacchi, 1836) * Mitromorpha olivoidea (Cantraine, 1835) Notes. The nomenclature of the two species belonging to the genus Mitromorpha , previously reported in Fumanti (2014) are here updated ac- cording to Amati et al. (2015). Ordo HETEROSTROPHAP. Fischer, 1885 Familia OMALOGYRIDAE G.O. Sars, 1878 Genus Omalogyra Jeffreys, 1859 Omalogyra atomus (Philippi, 1841) Genus Ammonicera Vayssiere, 1893 Ammonicera cfr. andresi Oliver et Rolan, 2015 * Ammonicera cW.fisclieriana (Monterosato, 1869) Ammonicera cfr. superstriata Oliver et Rolan, 2015 Notes. The determination of the species belong- ing to the genus Ammonicera , according to the recent review of this genus (Oliver & Rolan, 2015) and without SEM observations, has led to consid- erable difficulties and is proposed here with a wide margin of uncertainty. Familia PYRAMIDELLIDAE Gray, 1840 Genus Parthenina Bucquoy, Dautzenberg et Dollfiis, 1883 Parthenina clathrata (Jeffreys, 1848) * Parthenina dollfusi (Kobelt, 1903) * Parthenina emaciata (Brusina, 1866) * Parthenina interstincta (J. Adams, 1797) * Parthenina monozona (Brusina, 1869) *Partenina moolenbeechi (Amati, 1987) *Partenina penchynati (Bucquoy, Dautzen- berg et Dollfiis, 1883) Partenina suturalis (Philippi, 1844) Notes. As regards to Pyramidellidae we decided to include the full list of the genera and species The molluscan thanatocoenosis of Zannone Island (Pontine Archipelago, Latium, Italy). Additional reports 91 reported for Zannone (both in this paper and in Fumanti, 2014) with the nomenclature updated according to Giannuzzi-Savelli et al. (2014). Genus Folinella Dali et Bartsch, 1904 *Folinella excavata (Philippi, 1844) Genus Odostomella Bucquoy, Dautzenberg et Dollfus, 1883 * Odostomella doliolum (Philippi, 1844) Odostomella bicincta (Tiberi, 1868) Genus Euparthenia Thiele, 1931 *Euparthenia humboldti (Risso, 1826) Genus Eulimella Forbes et Mac Andrew, 1846 Eulimella acicula (Philippi, 1836) *Eulimella ventricosa (Forbes, 1844) Genus Odostomia Fleming, 1813 *Odostomia carrozzai Van Aartsen, 1987 *Odostomia eulimoides Hanley, 1844 * Odostomia lukisii Jeffreys, 1859 * Odostomia scalaris Mac Gillivray, 1843 Odostomia striolata (Forbes etHanlay, 1850) * Odostomia turrita Hanley, 1844 * Odostomia unidentata (Montagu, 1803) Genus Megastomia Monterosato, 1884 Megastomia alungata (Nordsieck, 1972) * Megastomia conoidea (Brocchi, 1814) Genus Ondina De Folin, 1870 *Ondina vitrea (Brusina, 1866) Ondina scadens (Monterosato, 1844) Genus Pyrgostylus Monterosato, 1884 *Pyrgostylus striatulus (Linnaeus, 1758) Genus Turbonilla Risso, 1826 *Turbonilla pumila G. Seguenza, 1876 Genus Careliopsis Morch, 1875 Careliopsis modesta (De Folin, 1870) Familia MURCHISONELLIDAE Casey, 1904 Genus Ebala Gray, 1 847 Ebala pointeli (De Folin, 1867) Familia CIMIDAE Waren, 1993 Genus Cima Chaster, 1896 Cima cylindrica (Jeffreys, 1856) Cima minima (Jeffreys, 1858) Familia TOFANELLIDAE Bandel, 1995 Genus Graphis Jeffreys, 1867 Graphys albida (Kanmacher, 1798) Ordo CEPHALAPSIDEA Fischer, 1883 Familia PLEUROBRANCHIDAE Gray, 1827 Genus Berthella Blainville, 1 824 Berthella sp. Familia RETUSIDAE Thiele, 1925 Genus Volvulella Newton, 1891 Volvulella acuminata (Broguiere, 1792) Familia PHIL1NIDAE Gray, 1850 Genus Philine Ascanius, 1772 Philine catena (Montagu, 1803) Philine angulata Jeffreys, 1867 Genus Petalifera Gray, 1 847 Petalifera cf. gravieri (Vayssiere, 1906) Familia AGLAJIDAE Pilsbry, 1895 Genus Spinoaglaja Ortea, Moro et Espinosa, 2007 Spinoaglaja wilpretii (Ortea, Bacallado et Noro, 2003) Notes. Species reported on the basis of one specimen (3.3 mm) devoid of soft part at 36 m of depth. 92 Bruno Fumanti & Italo Nofroni Classis BIVALVIA Linnaeus, 1758 Ordo SOLEMYOIDA Dali, 1889 Familia NUCULIDAE Gray, 1824 Genus Nucula Lamarck, 1799 Nucula sp. (juv.) Ordo MYTILOIDA Ferussac, 1822 Familia MYTILIDAE Rafmesque, 1815 Genus Crenella T. Brown, 1 827 Crenella pellucida (Jeffreys, 1850) Genus Dacrydium Torell, 1859 Dacrydium hyalinum Monterosato, 1875 Familia ANOMIIDAE Rafmesque, 1815 Genus Pododesmus Philippi, 1837 Pododesmus sp. Ordo LUCINIDAE Gray, 1854 Familia LUCINIDAE Fleming, 1828 Genus Anodontia Link, 1807 Anodontia fragilis Philippi, 1836 Genus Loripes Poli, 1791 Loripes lucinalis (Lamarck, 1818) Familia MONTACUTIDAE W. Clark, 1855 Genus Montacuta Turton, 1 822 Montacuta substriata (Montagu, 1808) Ordo VENEROID A Gray, 1854 Familia CARDIIDAE Lamarck, 1809 Genus Laevicardium Swainson, 1 846 Laevicardium crassum (Gmelin, 1791) Familia TELLINIDAE Blainville, 1814 Genus Arcopagia Brown, 1877 Arcopagia balaustina (Linnaeus, 1758) Familia PSAMMOBIIDAE Fleming, 1828 Genus Gari Schumacher, 1817 Gari costulata Turton, 1822 Gari depress a (Pennant, 1777) Classis SCAPHOPODA Broun, 1862 Ordo DENTALIIDAE da Costa, 1776 Familia DENTAL ID AE J.E. Gray, 1834 Genus Antalis H. Adams et A. Adams, 1854 Antalis vulgaris da Costa, 1778 REFERENCES Amati B., Smriglio C. & Oliverio M., 2015. Revision of the recent mediterranean species of Mitromorpha Carpenter, 1865 (Gastropoda, Conoidea, Mitro- morphidae) with the descriptions of seven new species. Zootaxa, 293 1 : 151-195. Fumanti B., 2014. Contribution to the knowledge of benthic molluscan thanatocoenosis of Zannone Island (Pontine Archipelago, Latium, Italy). Biodiversity Journal, 5: 97-106. Giannuzzi-Savelli R., Pusateri F., Micali P., Nofroni I. & Bartolini S., 2014. Atlante delle conchiglie marine del Mediterraneo. Vol. 5. Heterobranchia. Edizioni Danaus, Palermo. Ill + 91 pp. Oliver J.D. & Rolan E., 2015. The genus Ammonicera (Heterobranchia, Omalogyridae) in the Eastern Atlantic, 1 : the species of Iberian Peninsula. Iberus, 33: 45-95. Romani L. & Pagli A., 2014. The Genus Spinoaglaja Ortea, Moro et Espinosa, 2007 in the Mediterranean Sea: new records and observations on shell variability (Opisthobranchia, Aglajidae). Bollettino Malacolo- gico, 50: 137-139. Biodiversity Journal, 2016, 7 (1): 93-102 Monograph Terrestrial gastropods (Mollusca Gastropoda) from Lepini Mountains (Latium, Italy): a first contribution Alessandro Hallgass 1 & Angelo Van nozzi 2 'Via della Divina Provvidenza 16, 00166 Rome, Italy; e-mail: hallgass@hotmail.com 2 Via M.L. Longo 8, 00151 Rome, Italy; e-mail: ang.vannozzi@gmail.com ABSTRACT Lepini Mountains are a calcareous massif that forms the southern pre- Apennines of Latium (Italy), reaching a maximum altitude of 1536 m. Notwithstanding their central position and the low height reached, the malacofauna of Lepini Mountains has been long neglected and species composition was never reported so far. In this contribution, a preliminary investigation of the terrestrial gastropods (Mollusca Gastropoda) occurring in the Lepini Mountains is reported. At least 43 species are recorded. Several species already reported from Central Apen- nines occur. The most remarkable findings include a hitherto unrecorded population of Medora sp. (Clausiliidae) and the occurrence of two distinct forms ascribable to Jaminia quadridens s.l. KEY WORDS Terrestrial gastropods; biodiversity; Lepini Mountains; Italy. Received 28.02.2016; accepted 20.03.2016; printed 30.03.2016 Proceedings of the Ninth Malacological Pontine Meeting, October 3rd-4th, 2015 - San Felice Circeo, Italy INTRODUCTION Lepini Mountains, together with Aurunci and Ausoni Mountains, form the southern pre-Apen- nines of Latium (Italy). They are positioned about 50 km SE of Rome and extend in NE-SW direction (Fig. 1). They are separated from Central Apennines by the Sacco Valley and face the Pontine alluvial plain in the south. Lepini Mountains are comprised of two parallel chains directed in NE-SW direction, separated by the deep Montelanico-Carpineto- Maenza tectonic line. The western chain is com- prised by Mount Lupone (1378 m) and Mount Semprevisa group (1536 m, the highest peak), whereas the eastern chain quickly slopes down to the Sacco Valley and is comprised of Mount Gemma, Mount Malaina, Mount S. Marino and Mount Alto, all of which reach heights around 1400 m. Lepini Mountains are mainly comprised of limestone of Cretaceous age (Sani et al., 2004). The whole massif shows to intense karst phenomena. As a consequence, in the Lepini Mountains no per- manent water body occurs, whereas several springs appear at the base of the massif, the best known of which gives rise to the Oasis of Ninfa (Amori et al., 2002 ). The vegetation is mainly comprised of holm oak, chestnut and mixed woods at medium-low altitudes and beech forests at medium-high alti- tudes. Large portions of territory are occupied by grassland mainly used as pasture. The invertebrate fauna of Lepini Mountains have been studied in some detail with regard to arthropods (Corsetti et al., 2015). Several studies focused on hypogean fauna (Sbordoni, 1971; Latella, 1995; Nardi et al., 2002). Moreover, some endemisms have been reported (Sbordoni, 1971; 94 Alessandro Hallgass & Angelo Vannozzi Figure 1. Studied area: Lepini Mountains (southern Latium, Italy). Numbers indicate the sampled stations listed in Table 1 . Stn. Locality Coordinates Alt. (m) Environment 1 Pass to Campo di Segni 41.672649° N, 12.987930° E 1015 Pasture with stones 2 After pass to Campo di Segni 41.670720° N, 12.993717° E 960 Rocks with low vegetation 3 Close to Campo di Segni 41.667926° N, 12.990029° E 880 Pasture and bushes wit stones 4 Mount Erdigheta 41.56410° N, 13.119629° E 1046 Beech forest with rocks 5 Mount Erdigheta 41.562092° N, 13.120613° E 1115 Pasture with stones 6 Mount Semprevisa 41.572228° N, 13.092678° E 1250- 1400 Beech forest with rocks 7 Mount Semprevisa, top 41.571147° N, 13.091063° E 1490 Stones on the top 8 Carpineto, Pian della Faggeta 41.575702° N, 13.103665° E 930 Rocks with low vegetation and residuary beeches 9 Bassiano, road to Sempre- visa, near the spring 41.552975° N, 13.047529° E 590 Holm oak wood with rocks 10 Bassiano, road to Semprevisa 41.558473° N, 13.059121° E 864 Clearing in holm oak wood 11 Campo Rosello 41.563711° N, 13.077542° E 1174 Pasture with stones 12 Campo Rosello 41.571987° N, 13.072649° E 41.574551° N, 13.074571° E 1250- 1410 Pasture with stones Table 1. List of the stations: Lepini Mountains (southern Latium, Italy). Terrestrial gastropods (Mollusca Gastropoda) from Lepini Mountains (Latium, Italy): a first contribution 95 Pace, 1975; Magrini, 2005). However, the mol- luscan fauna of Lepini Mountains was never studied so far. Only four species occurring in hypogean environments were reported ( Discus rotundatus, Campylaea planospira, Daudebardia brevipes and Oxychilus draparnaudi ), none of which strictly hypogean (Latella, 1995). In this contribution, we report the results of a first survey aimed at assessing the biodiversity of terrestrial gastropods of Lepini Mountains. MATERIAL AND METHODS A total of 1 2 stations along the western chain have been sampled between April and June 2015 (see Table 1). Sampling was carried out only in natural environments. As a consequence, species recorded only from urban areas, such as Cornu aspersum (O.F. Muller, 1774) found in the town of Bassiano, were ruled out. Additionally, freshwater or hypogean environments were not considered. For the nomenclature, we mainly referred to the check- list of the species of the Italian fauna (Bodon et al., 1995; Manganelli et al., 1995, 1998, 2000). For the suprageneric nomenclature, we referred to Bouchet & Rocroi (2005). All specimens here illustrated were collected from the Lepini Mountains. Shell length and width were measured parallel and perpendicular to the axis of the shell, respectively, with calipers to the nearest 0.1 mm. RESULTS AND DISCUSSION At least 44 species of terrestrial gastropods occur in the Lepini Mountains (see Table 2). The most speciose family is the Clausiliidae, with 7 recorded species. The clausiliid Leucostigma can- didescens is by far the commonest and widespread species, occurring in almost all calcareous outcrops, either exposed or shaded, often associated with other calciophilous species such as Cochlostoma cf. adamii, Marmorana signata, Granaria apennina and Medora sp. We agree with Feher et al. (2010) who indicate th species of the genus Granaria Held, 1838 occurring in the Italian peninsula as G. apen- nina. Medora sp. was found only in a single station inside the beech forest of Mount Semprevisa. The unexpected finding of this population confirms that the current knowledge of the genus Medora H. et A. Adams, 1855 in Italy is far from being exhaust- ive (Giusti et al., 1986; Nordsieck, 2012; Colomba et al., 2012). Cochlodina laminata and C. bidens occur in sympatry in the beech forest. According to Opinion 2355, the Apennine species so far known as Cochlodina incisa (Kuster, 1876) should be indicated as C. bidens (Linnaeus, 1758) (Kadolsky, 2009; ICZN, 2015). They are readily distinguished by the development of palatal plicae. In fact, while in the former palatal plicae are truncated at the level of the clausilium, in the latter both the principal and the lower palatal plicae prolong internally. Moreover, an additional intermediate palatal plica often occurs in the latter. Cochlodina bidens in Lepini Mountains shows a stout shell also found in specimens from other localities of Latium, such as the holm oak woods of Mount Circeo and Macchia Grande (Fi- umicino) (Hallgass & Vannozzi, 2014). The populations of Cochlostoma cf. adamii have been studied by Zallot et al. (2015) in the generic revision of the family Cochlostomatidae and assigned to the subgenus Turritus Westerlund, 1883. Cochlostoma adamii group is comprised of several forms reported with different nominal taxa occurring from Central Apennines to Sicily, whose taxonomy needs to be clarified. The marquise Paulucci (1881) noted the occurrence of foims close to Pomatias adamii in the Central Apennines and described “ Pomatias adamii Var. Carseolanus“ from Carsoli (Abruzzi). Cochlostoma cf. adamii from Lepini Mountains is different from C. cassi- niacum (Saint Simon in Paulucci, 1878) from Cas- sino and Sterrone (both Latium), though belonging to the same subgenus Turritus (Zallot, comm, pers.). On the whole, the beech forest shows the greatest biodiversity, with 28 recorded species. Among them, there are several species commonly found in beech forests of Central Apennines (Giusti et al., 1985). However, a few of them deserve some comments. Acicula sp. was recorded from a worn fragment. The closest finding of this genus is A. szigethyannae Subai, 1977 from Val d’Arano (Ovindoli, Abruzzi). Conversely, Platyla similis is recorded from several localities of the Italian penin- sula. In particular, it has been reported from the neighbouring Aurunci Mountains (Bodon & Cianfanelli, 2008). Umax cf. maximus appears with different patterns (Figs. 8 and 9). A completely 96 Alessandro Hallgass & Angelo Vannozzi Family Species Stn. COCHLOSTOMATIDAE Cochlostoma cf. adamii (Paulucci, 1879) 4-9, 11, 12 2 ACICULIDAE Acicula sp. 6 11 Platyla similis (Reinhardt, 1880) 6 10 POMATIIDAE Pomatias elegans (O.F. Muller, 1774) 2,3,9 ORCUL1DAE Sphyradium doliolum (Bruguiere, 1792) 6, 8 VALLONIIDAE Acanthinula aculeata (O.F. Muller, 1774) 8,9 13 Gittenbergia sororcula (Benoit, 1857) 6 15 CHONDRINIDAE Granaria apenninci (Kuster, 1850) 4,8 Chondrina avenacea (Bruguiere, 1792) 6,9 VERTIGINIDAE Truncatellina callicratis (Scacchi, 1833) 9 14 ENIDAE Jaminia quadridens (O.F. Muller, 1774) (small morphotype) 7,8 5, 18 Jaminia quadridens (O.F. Muller, 1774) (large morphotype) 4, 5, 7, 8, 12 4, 19 Merdigera obscura (O.F. Muller, 1774) 6, 8 FERUSSACIIDAE Cecilioides acicula (O.F. Muller, 1774) 9 SUBULINIDAE Rumina decollata (Linnaeus, 1758) 1,3,9 CLAU SILIID AE Medora sp. 6 20 Leucostigma candidescens (Rossmassler, 1835) 1-6, 8, 9, 11, 12 3,21 Cochlodina laminata (Montagu, 1803) 6 23 Cochlodina bidens (Linnaeus, 1758) 6 22 Siciliaria paestana (Philippi, 1836) 3, 5, 6, 9 Macrogastra plicatula (Draparnaud, 1801) 6 Clausilia cruciata Studer, 1 820 6 PUNCTIDAE Punctum pygmaeum (Draparnaud, 1801) 6 12 DISCIDAE Discus rotundatus (O.F. Muller, 1774) 6,9 PRISTILOMATIDAE Vi Ire a botterii (Pfeiffer, 1853) 4, 6,8 17 Vitrea subrimata (Reinhardt, 1871) 6, 8 16 OXY CHIL1D AE Daudebardia rufa (Draparnaud, 1805) 6 6 Daudebardia brevipes (Draparnaud, 1805) 6 Oxy chilus cf. draparnaudi (Beck, 1837) 2, 6, 8,10 MILACIDAE Tandonia sowerbyi (Ferussac, 1823) 6, 10 7 VITRINIDAE Semilimacella bonellii (Targioni Tozzetti, 1873) 6 LIMACIDAE Umax cf. maximus Linnaeus, 1758 3, 6, 10 8,9 Umax sp. A (black) 3,6, 8 9 Umax sp. B (brown) 6 AGRIOLIMACIDAE Deroceras cf. lothari Giusti, 1973 8 H Y GROMIID AE Monacha cf. cantiana (Montagu, 1803) 9 Monacha cf. campanica (Paulucci, 1881) 1,3, 5, 8, 11, 12 Cernuellopsis ghisottii Manganelli et Giusti, 1988 1,4, 5,7, 8,11,12 Cernuella cisalpina (Rossmassler, 1837) 10 Hygromia cinctella (Draparnaud, 1801) 1 HELICIDAE Campylaea planospira Lamarck, 1 822 6 Marmorana signata (Ferussac, 1821) 6, 9, 11 Cantareus apertus (Bom, 1778) 9 Helix ligata O.F. Muller, 1774 1,3, 12 24 Table 2. List of the species recorded in the sampled stations, Lepini Mountains (southern Latium, Italy). Terrestrial gastropods (Mollusca Gastropoda) from Lepini Mountains (Latium, Italy): a first contribution 97 Figures 2-9. Terrestrial gastropods from Lepini Mountains. Fig. 2: Cochlostoma cf. adamii. Fig. 3: Leucostigma candides- cens. Fig. 4: Jaminia qnadridens (large morphotype). Fig. 5: Jaminia quadridens (small morphotype). Fig. 6: Daudebardia rufa. Fig. 7: Tandonia sowerbyi. Fig. 8: Umax cf. maximus. Fig. 9: Limax sp. A (black) and L. cf. maximus. 98 Alessandro Hallgass & Angelo Vannozzi Figures 10-17. Terrestrial gastropods from Lepini Mountains. Fig. 10: Platyla similis. Fig. 1 1 : Acicula sp. Fig. 12: Punctum pygmaeum. Fig. 13: Acanlhinula aculeata. Fig. 14: Truncatellina callicratis. Fig. 15: Gittenbergia sororcula. Fig. 16: Vitrea subrimata. Fig. 17: Vitrea botterii. Scale bar: 1 mm. Terrestrial gastropods (Mollusca Gastropoda) from Lepini Mountains (Latium, Italy): a first contribution 99 Figures 18-23. Terrestrial gastropods from Lepini Mountains. Fig. 18: Jaminia quadridens (small morphotype). Fig. 19: Jaminia quadridens (large morphotype). Fig. 20: Medora sp. Fig. 21: Leucostigma candidescens . Fig. 22: Cochlodina bidens. Fig. 23: Cochlodina laminata. Scale bar: 5 mm (whole specimens); 2.5 mm (details). 100 Alessandro Hallgass & Angelo Vannozzi black specimens of Limax Linnaeus, 1758 was found in different stations, also in sympatiy with L. cf. maximus (Fig. 9). A further Limax species with uniform brown colour was recorded. A small black specimen of Deroceras Rafinesque, 1820 was collected in Stn. 8, externally resembling Deroceras lothari, a species described as endemic to Reatini Mountains (northen Latium) (Giusti, 1973). It is provisionally reported as Deroceras cf. lothari pending further study. Holm oak and mixed woods are poorer in terms of both species and number of specimens. Pastures show a relatively reduced number of species as well, some of which deserve some comments. Jaminia quadridens occurs in two distinct forms, here referred to as small and large morphotype, respectively (Figs. 4, 5, 19, 20). At a conchological level, they mainly differ with regard to shell size. Measurements on over 60 specimens show that these two forms can be readily distinguished by the shell width, which shows a clear gap between the two morphotypes (Fig. 25). Moreover, the large morphotype seems to show a proportionally larger width. However, this feature needs confirmation due to the small amount of recorded specimens of the small morphotype. The large morphotype is widespread and was recorded from several stations. Conversely, the small morphotype is uncommon and was recorded only at higher altitudes. A similar scenario with different morphs of J. quadridens occurring syntopically have been recorded also in other localities of Central Apennines, often with the occurrence of an additional, dextral morph. Monacha campanica, which we regard as a dis- tinct species, is found throughout the Liri Valley up to high altitude. The genitalia of M. campanica are characterized by a very short penis provided with a large penial papilla and a slender epiphallus. The flagellum is as long as the epiphallus. The bursa copulatrix shows a thick duct, widened at the base. Specimens from Lepini Mountains share the same anatomy but show a rather different shell. In fact, they are smaller (max. diam. 16 mm) and lighter than those described by the marquise Paulucci (1881) (max. diam. 21 mm). Moreover, they show a less depressed spire and a narrower umbilicus. As a consequence, they are reported in Table 2 as Monacha cf. campanica , pending further study. In Stn. 9 (Bassiano, road to Semprevisa, near the spring) an empty shell clearly different from Figure 24. Helix ligata, height 35 mm, with genitalia (scale bar: 5 mm). Terrestrial gastropods (Mollusca Gastropoda) from Lepini Mountains (Latium, Italy): a first contribution 101 Jaminia quadridens s.l. Fig. 25. Height and width measurements of Jaminia quad- ridens s.l. specimens from Lepini Mountains. Best-fit lines for the two morphotypes are shown. Monaca cf. campanica has been collected. It is provisionally reported as Monacha cf. cantiana waiting for anatomical data. Cernuellopsis ghisottii is the most abundant species in grasslands and pastures above 1000 m. This species shows a disjunct distribution in the Apennines. Southern populations occur in the Pollino (Calabria-Basilicata) and Sirino (Basilicata) massifs and extend up to Albumi Mountains (Cam- pania), wherease populations from Central Apen- nines are frequent on the western coastal chains and extend up to Simbruini Mountains (Latium). The species was never recorded from intermediate mountains of central-northen Campania. Helix ligata is widespread in Lepini Mountains but never abundant. Different populations show a variable appearance. Specimens from Carpinetana Valley were genetically studied by Fiorentino et al. (2016) and are very similar to specimens from Apennine valley floors showing a yellowish background due to the presence of periostracum, whereas specimens from Stn. 1 (pass to Campo di Segni) living in exposed pasture with stones show a whitish background (Fig. 24) likely due to the loss of periostracum that make it resemble Helix delpre- tiana Paulucci, 1878 (Giusti, 1973). However, the anatomy of specimens from Stn. 1 corresponds to Helix ligata , even though they are genetically dis- tinct from populations from Central Apennines (Fiorentino et al., 2016). CONCLUSIONS The preliminary checklist here presented shows that at least 43 species of terrestrial gastropods occur in the Lepini Mountains. The richest environ- ment is represented by beech forests, with 28 recor- ded species. Along with species already reported from Central Apennines, a new isolated population of Medora is recorded. Due to the extremely limited distribution, this population can be considered vul- nerable and likely in need of protection. Jaminia quadridens occurs in two clearly distinct morphs without intermediate forms, mainly differing in their size. Further research is required in order to ascertain whether they actually belong to different species. Some species remain undetermined, whereas others were determined by comparison, pending further research. REFERENCES Amori G., Corsetti L. & Esposito C, 2002. Mammiferi dei Monti Lepini. Quaderni di Conservazione della Natura, 1 1 , Ministero dell'Ambiente e della Tutela del Territorio - Istituto Nazionale per la Fauna Selvatica "Alessandro Ghigi". Bodon M. & Cianfanelli S., 2008. Una nuova specie di Platyla per il sud Italia (Gastropoda: Prosobranchia: Aciculidae). Bollettino Malacologico, 44: 27-37. Bodon M., Favilli L., Giannuzzi Savelli R., Giovine F., Giusti F., Manganelli G., Melone G., Oliverio M., Sabelli B. & Spada G., 1995. Gastropoda Proso- branchia, Heterobranchia Heterostropha. In: Minelli A., Ruffo S. & La Posta S. (Eds), Checklist delle specie della fauna italiana 14, Calderini, Bologna, 60 pp. Bouchet Ph. & Rocroi J.P., 2005. Classification and nomenclator of gastropod families. Malacologia, 47: 1-397. Colomba M.S., Liberto F., Reitano A., Renda W., Pocaterra G., Gregorini A. & Sparacio I., 2012. Molecular studies on the genus Medora H. et A. Adams, 1855 from Italy (Gastropoda Pulmonata Clausiliidae). Biodiversity Journal, 3: 571-582. Corsetti L., Angelini C., Copiz R., Mattoccia M. & Nardi G., 2015. Biodiversita dei Monti Lepini. Edizioni Belvedere, Latina, 352 pp. 102 Alessandro Hallgass & Angelo Vannozzi Feher Z., Deli T. & Solymos P., 2010. Revision of Granaria frumentum (Draparnaud 1801) (Mollusca, Gastropoda, Chondrinidae) subspecies occurring in the eastern part of the species’ range. Journal of Conchology, 40: 201-217. Fiorentino V., Manganelli G., Giusti F. & Ketmaier V., 2016. Recent expansion and relic survival: Phylogeo- graphy of the land snail genus Helix (Mollusca, Gastropoda) from south to north Europe. Molecular Phylogenetics and Evolution, 98: 358-372. Giusti F., 1973. Notulae malacologicae XVI. I molluschi terrestri e di acqua dolce viventi sul massiccio dei Monti Reatini (Appennino Centrale). Lavori della Societa Italiana di Biogeografia, NS 2: 423-574. Giusti F., Castagnolo L. & Manganelli G., 1985. La fauna malacologica delle faggete italiane: brevi cenni di ecologia, elenco delle specie e chiavi di riconosci- mento dei generi e delle entita piu comuni. Bollettino Malacologico, 21: 69-144. Giusti F., Grappelli C., Manganelli G., Fondi R. & Bulli L., 1986. An attempt of natural classification of the genus Medora in Italy and Yugoslavia, on the basis of conchological, anatomical and allozymic charac- ters (Pulmonata: Clausiliidae). Lavori della Societa Italiana di Malacologia, 22: 259-341. Hallgass A. & Vannozzi A., 2014. The continental molluscs from Mount Circeo (Latium, Italy). Biod- iversity Journal, 5: 151-164. ICZN, 2015. Opinion 2355 (Case 3581): Turbo bidens Linnaeus, 1758 (Gastropoda, Clausiliidae): request to set aside the neotype not granted. Bulletin of Zoological Nomenclature, 72: 159-161. Kadolsky D., 2009. Turbo bidens Linnaeus 1758 (Gastro- poda: Clausiliidae) misidentified for 250 years. Journal of Conchology, 40: 19-30. Latella L., 1995. La fauna cavernicola dei Monti Lepini. Notiziario del Circolo Speleologico Romano, NS 6- 7, 1991-'92: 78-119. Magrini P., 2005. Un nuovo Neobacanius anoftalmo del Lazio (Insecta, Coleoptera: Histeridae). Aldrovandia, 1: 55-62. Manganelli G., Bodon M., Favilli L. & Giusti F., 1995. Gastropoda Pulmonata. In: Minelli A., Ruffo S. & La Posta S. (Eds.), Checklist delle specie della fauna italiana 16, Bologna (Calderini): 1-60. Manganelli G., Bodon M., Favilli L., Castagnolo L. & Giusti F., 1998. Checklist delle specie della fauna italiana, molluschi terrestri e d’acqua dolce. Errata e addenda. Bollettino Malacologico, 33: 151-156. Manganelli G., Bodon M. & Giusti F., 2000. Checklist delle specie della fauna italiana, molluschi terrestri e d’acqua dolce. Errata e addenda. Bollettino Malaco- logico, 36: 125-130. Nardi G., Di Russo C. & Latella L., 2002. Populations of Nepa cinerea (Heteroptera: Nepidae) from hy- pogean sulfurous water in the Lepini Mountains (Latium, Central Italy). Entomological News, 113: 125-130. Nordsieck H., 2012. Erganzung der Revision der Gattung Medora H. & A. Adams: Die Medora- Alien Italiens (Gastropoda, Stylommatophora, Clausiliidae, Alopi- inae), mit Beschreibung einer neuen Unterart von Medora dalmatina Rossmassler. Conchylia, 42: 75- 81. Pace R., 1975. An exceptional endogeous beetle: Crowsoniella relicta n. gen. n. sp. of Archostemata Tetraphaleridae from central Italy. Bollettino del Museo Civico di Storia Naturale di Verona, 2: 445- 458. Paulucci M., 1881. Contribuzione alia fauna malacolo- gica italiana. Specie raccolte dal Dott. C. Cavanna negli anni 1878, 1879, 1880 con elenco delle conchiglie abruzzesi e descrizione di due nuove Succinea. Bullettino della Societa Malacogica Italiana, 7: 69-180. Sani F., Del Ventisette C., Montanari D., Coli M., Nafissi P. & Piazzini A., 2004. Tectonic evolution of the internal sector of the Central Apennines, Italy. Marine and Petroleum Geology, 21: 1235-1254. Sbordoni V., 1971. II popolamento animale e vegetale dell ’Appennino Centrale. Lavori della Societa Italiana di Biogeografia, NS 2: 595-614. Zallot E., Groenenberg D.S.J., De Mattia W., Feher Z. & Gittenberger E., 2015. Genera, subgenera and species of the Cochlostomatidae (Gastropoda, Caeno- gastropoda, Cochlostomatidae). Basteria 78: 63-88. Biodiversity Journal, 2016, 7 (1): 103-115 Monograph A revision of the Mediterranean Raphitomidae, 3: on the Raphitoma pupoides (Monterosato, 1 884) complex, with the description of a new species (Mollusca Gastropoda) Francesco Pusateri 1 , Riccardo Giannuzzi-Savelli 2 * & Stefano Bartolini 3 'Via Castellana 64, 90135 Palermo, Italy; e-mail: francesco@pusateri.it 2 Via Mater Dolorosa 54, 90146 Palermo, Italy; e-mail: malakos@tin.it 3 Via E. Zacconi 16, 50137 Firenze, Italy; e-mail: stefmaria.bartolini@libero.it ’Corresponding author ABSTRACT In the present work we present a complex of species of the family Raphitomidae (Mollusca Gastropoda) comprising three entities: two have multispiral protoconchs, Raphitoma pupoides (Monterosato, 1884), the less known R. radula (Monterosato, 1884) and a new species with paucispiral protoconch. KEY WORDS Mollusca; Conoidea; Raphitomidae; new species; Mediterranean Sea. Received 02.03.2016; accepted 24.03.2016; printed 30.03.2016 Proceedings of the Ninth Malacological Pontine Meeting, October 3rd-4th, 2015 - San Felice Circeo, Italy INTRODUCTION The family of Raphitomidae is a well supported clade of the Conoidea (Bouchet et al., 2011). The genus Raphitoma Bellardi, 1847 as currently conceived includes, based on our estimates, ca. 40 Mediterranean species, some of which are still undescribed. Propaedeutic to the general revision of the Mediterranean Raphitoma s.l., we have focused on several pairs of species, differing only or mostly in the size and shape of the protoconch (Pusateri et al., 2012, 2013). The specific distinction is based on the assumption that the dichotomy multispiral protoconch/planktrotrophic develop- ment vs. paucispiral protoconch/lecithotrophic de- velopment (Jablonski & Lutz, 1980) can be used in caenogastropods to recognise distinct sister species (Bouchet, 1989; Oliverio, 1996a, 1996b, 1997). Anyway, it should not be abused to create poly- phyletic genera by artificially separating closely related species among different genera only based on their larval development (Bouchet, 1990). In the present work we present the results on a complex of species comprising three entities: two have multispiral protoconchs, R. pupoides (Monterosato, 1884), and the less known R. radula (Monterosato, 1884); the other was discovered while revising the materials in the Monterosato collection, where a lot (MCZR 16905) included some specimens with paucispiral protoconch, la- belled by Monterosato himself “K tomentosa/ Monts. /Palermo’', never published, that we describe hereby as new to Science. ABBREVIATIONS AND ACRONYMS, d = diameter; h = height; sh = empty shell(s); LMG-NS: Leeds Museums and Galleries - Natural Science; MNHN: Musee Nationale Histoire Naturelle, Paris, France; MRSNT: Museo Regionale Storia Naturale, Terras ini, Italy; NMW: National Museum of Wales, United Kingdom; SMF: Senckenberg Museum, 104 Francesco Pusateri etalii Frankfurt/M, Germany; SMNH: Swedish Museum of Natural History, Stockholm, Sweden; MCZR: Museo Civico di Zoologia, Roma, Italy; HUJ: Hebrew University of Jerusalem, Israel; ARD: Roberto Ardovini collection (Rome, Italy); BOG: Cesare Bogi collection (Livorno, Italy); DUR: Sergio Duraccio collection (Napoli, Italy); GER: Alfio Germana collection (Trecastagni, Catania, Italy); GOR: Sandro Gori collection (Livorno, Italy); HOA: Andre Hoarau collection (Frejus, France); MAC: Gabriele Macri collection (Scor- rano, Lecce, Italy); MAR: Alessandro Margelli collection (Livorno, Italy); PAG: Attilio Pagli collection (Lari, Pisa, Italy); PAO: Paolo Paolini collection (Livorno, Italy); PRK: Jakov Prlcic collection (Split, Croatia); PSI: Peter Sossi collec- tion (Trieste, Italy); PUS: Francesco Pusateri col- lection (Palermo, Italy); SBR: Carlo Sbrana collection (Livorno, Italy); SER: Gabriele Sercia collection (Palermo, Italy); SPA: Gianni Spada collection (Vagrigneuse, France); SQU: Ennio Squizzato collection (Loreggia, Padova, Italy); TIS: Morena Tisselli collection (S. Zaccaria, Ravenna, Italy); TRI: Lionello Tringali collection (Rome, Italy); VAZ: Angelo Vazzana collection (Reggio Calabria, Italy). RESULTS Systematic Citation of unpublished names is not intended for taxonomic purposes. Familia RAPHITOMIDAE Bellardi, 1875 Genus Raphitoma Bellardi, 1 847 Type species: Pleurotoma hystrix Cristofori et Jan, 1832 (nomen nudum, validated by Bellardi, 1847 as " Pleurotoma histrix Jan.") by subsequent des- ignation (Monterosato, 1872: 54). Raphitoma pupoides (Monterosato, 1884) Figs. 1-9, 24 Pleurotoma rudis Scacchi, 1836 non G.B. Sowerby I, 1834 nee Philippi, 1836 Pleurotoma rudis Scacchi, Weinkauff, 1868: 130 (see Remarks) Pleurotoma reticulatum var. rudis Sc., Petit de la Saussaye, 1869: 154 Pleurotoma ( Defrancia ) rudis Sc., Monterosato, 1875: 44 (see Remarks) Pleur. rude Scacchi, Aradas & Benoit, 1876: 249 n. 662 (see Remarks) Pleurotoma rudis Sc., Monterosato, 1878: 106 (see Remarks) Clathurella rudis Scacchi, B.D.D., 1883: 94 pi. 14 figs. 8, 9 Cordieria pupoides Monterosato, 1884: 132 [nomen novum] Clathurella pupoidea Monterosato, Locard, 1886: 114 [error pro pupoides] Clathurella pupoidea Monterosato, Locard, 1891: 66 fig. 52 [error pro pupoides] Clathurella rudis (B.D.D.), Cams, 1893: 426 Clathurella pupoidea de Monterosato, Locard & Caziot, 1900: 248 Clathurella pupoidea var. major, Locard & Caziot, 1900: 248 (nomen nudum) Clathurella pupoidea var. minor, Locard & Caziot, 1900: 248 (nomen nudum) Clathurella pupoidea var. ventricosa, Locard & Caziot, 1900: 248 (nomen nudum) Clathurella pupoidea var. curta, Locard & Caziot, 1900: 248 (nomen nudum) Clathurella pupoidea Mtrs., Kobelt, 1905: 351 Mangilia ( Clathurella ) pupoides Monterosato, Cipolla, 1914: 146, pi. 13, figs. 16 (fossil)-17 (recent) Cordieria pupoides Montrs., Bellini, 1929: 32 Philbertia ( Philbertia ) rudis Scacchi, Priolo, 1967: 697 Raphitoma ( Cyrtoides ) rudis (Scacchi), Nordsieck, 1968: 176 pi. 30, fig. 20 Raphitoma ( Cyrtoides ) rudis pupoidea (Monterosato), Nordsieck, 1968: 176 pi. 30 fig. 21 Raphitoma rudis pupoidea Monts, Parenzan, 1970: 207 pi. 44, fig. 842 Raphitoma (C.) pupoidea (Monterosato), Nordsieck, 1977: 52, pi. 16, fig. 126 (error pro pupoides ) Raphitoma (C.) neapolitana Nordsieck, 1977: 52, pi. 16 figs. 124, 125 (nomen vanum) Raphitoma pupoides (Mts), Terreni, 1981 : 40 n. 328 Raphitoma pupoidea (Monterosato), Nordsieck, 1982: 272, pi. 101, fig. 98.11 Raphitoma neapolitana Nordsieck, 1982: 272, pi. 101, fig. 98.10 Raphitoma neapolitana form a Nordsieck, 1982: 272, pi. 101, fig. 98.10a Mediterranean Raphitomidae, 3: on the Raphitoma pupoides complex, with the description of a new species 105 Raphitoma (R. ) pupoides (Monterosato), Van Aartsen etal., 1984: 91 Raphitoma pupoides (Monterosato), Orlando & Palazzi, 1986: 44 Raphitoma pupoides (Monterosato), Tenekidis, 1989: n. 58.50 Raphitoma {Raphitoma) pupoides (Monterosato), Sabelli et al., 1990-1992: 44, 216, 411 Raphitoma pupoides (Monterosato), Poppe & Goto, 1991: 174 Raphitoma ( Cyrtoides ) pupoides (Monterosato), Delamotte & Vardala-Theodorou, 1994: 287 Raphitoma pupoides (Monterosato), Cecalupo & Quadri, 1995: 109 Raphitoma pupoides (Monterosato), Giribet & Penas, 1997: 53 Raphitoma pupoides (Monterosato), Marquet, 1998: 276 Raphitoma pupoides (Monterosato), Oztiirk et al., 2004: 59 Raphitoma pupoides (Monterosato), Repetto et al., 2005:220% 910 Pleurotoma rudis Scacchi, Cretella et al., 2005: 125 Raphitoma pupoides (Monterosato), Cretella et al., 2005: 125 Raphitoma pupoides (Monterosato), Cossignani & Ardovini, 2011: 31, 328 Raphitoma pupoides (Monterosato, 1884), Scuderi & Terlizzi, 2012 (see Remarks) Type locality. Coast of Provence, France, Mediterranean Sea. Examined material. Type material: neotype, from “Artufel/Provenza” [Provence, M. Artufel legit] (18.7 x 7.7 mm) (MCZR 16492). Other examined material. France. “Artufel/ Provenza” 3 sh (MCZR 16492, with Monts label “H. pupoides”); Marseille, 4 sh (coll. Locard MNHN); St. Raphael, 3 sh coll. Locard (MNHN), 1 sh (coll. Hoarau); Cassis, 2 sh coll. (Locard MNFIN); Le Brusc, 4 sh (coll. Locard MNHN, 4 sh); Coste di Provenza, 2 sh (coll. Chaster NMW n. 01894); Bastia, 2 sh (coll. Monterosato, MCZR lot 16861). Italy. Gulf of Baratti, 7 sh (PAO), 1 sh (PAG); Punta Ripalti (Elba Isl.), 2 sh -25 m (GOR); Lazio 1 sh (PAG); Circeo, 1 sh (TRI); Napoli, 1 sh (coll. Coen HUJ, n. 8082c sub nomine “ Philbertia ( Cordieria ) cordieri cancellatd’y Sorrento (Napoli), 2 sh (DUR); Palinuro (Salerno), 1 sh (SPA); Scilla (Reggio Calabria), 4 sh (VAZ); Palermo, Sicily, 10 sh with Monterosato handwritten label “pupoides /Monts. /ValU/et/v. decolorata, Pallary”, 1 sh with non-Monterosato label “Cordieria/ pup- oides Monts./dr. Golfo di Palermo” and 15 sh with non-Monterosato label “Cordieria/ pupoides Monts./ drag. Golfo di Palermo” (MCZR 16492, with Monts label “H. pupoides ”); Porticello (Palermo), 2 sh sub nomine R. reticulata (coll. MRSNT n. 4759); Isola delle Femmine (Palermo), 1 sh (SER), 8 sh (PUS); Trapani, 1 sh (SER); Catania, 1 sh (GER); Pozzillo Inferiore (Catania), 1 sh (PAG); Canale di Sicilia, 1 sh (TRI), 1 sh (coll. MRSNT n. 7312); Sicilia, 6 sh sub nomine R. purpurea (coll. MRSNT n. 29824); Jesolo (Venezia), 1 sh (SQU). “Coste d'Africa”. 1 sh, coll. Monterosato MCZR, lot 16901. Croatia. Unprecised locality, 1 sh (DEL); Dal- matia, 1 sh (PRK). Description. In squared parentheses data of the neotype. Shell of medium size for the genus, height 10-21 mm [18.7] (mean 15.05, std 3.81), width 5- 8 mm [7.7] (mean 6.57, std 1.27), cirto-pupoid, slender, h/d 2.1-2.57 [2.43] (mean 2.26, std 0.19). Protoconch multispiral, only part of the last whorl known, with traces of diagonally cancellate sculp- ture. Teleoconch of 6-8 [7] whorls, evenly convex (more convex in juveniles). Suture fine and undu- late. Axial sculpture of 12-24 [18] sligthly opistho- cline, non-equidistant ribs, and interspaces broader than the ribs (with interspace width varying with shell size). Axial sculpture evident, but becoming obsolescent in largest shells. In particularly large shells (gerontic), axial ribs revert to same strength as the spiral cords on the last quarter of whorl. Spiral sculpture on the last whorl of 7-10 [9] cordlets, thin- ner that axial ribs. Cancellation squared in juveniles, becoming rectangular in adults. Secondary cordlets appearing occasionally and thereafter becoming as strong as the others. Subsutural ramp narrow, devoid of evident sculpture. Columella simple, slightly sinuous anteriorly, gently angled posteriorly. Outer lip thickened and crenulated externally, with 11-13 [12] strong inner denticles, the most posterior smal- ler, delimiting the wide and short anal sinus, the most anterior more robust and delimiting the funnel- like siphonal canal. Siphonal fasciole of 6 nodulose cordlets, neatly spaced from the last spiral cordlet. Colour uniformly ligth chestnut brown in the back- 106 Francesco Pusateri etalii ground, with darker blotches, more evident in larger shells (>20 mm), and same darker colour bordering the siphonal fasciole and inside the aperture. Violet hue on the first 3^4 whorls of particularly fresh spe- cimens. Comma- shaped white spots on the sub- sutural ramp, arrow-like white spots inside some cancellation interspaces. Soft parts unknown. Distribution. Western and Central Mediter- ranean. Adriatic. The records under this name from Greece by Koukouras (2010) and Delamotte & Vardala-Theodorou (1994: 287) were in turn based on Tenekides (1989) who reported under this name another species (probably R. echinata ). Remarks. The protoconch was always either lacking, broken or corroded in almost specimens studied. Anyway parts of the apical whorls showing traces of a diagonally cancellate sculpture, indic- ating a multispiral protoconch. Pleurotoma rudis Scacchi, 1836 was introduced with the following diagnosis: “Testa fusca fascis pallidioribus, anfractibus rotundatis, cancellatis et muricatis; labro crasso interne striato, cauda vix ultra labrum producta. Alta lin. 10—11. R echinatae similis; at labro crassiore, cauda breviore, et minus aspera; saepe fascis pallidioribus ornata. In sinu neapolitano et tarentino ” (Scacchi, 1836), Fig. 17. Weinkauff (1868), Petit de la Saussaye (1869) and Aradas & Benoit (1876) considered it as a variety or synonym of R. echinata (as Defrancia reticulata Renier). Monterosato (1875, 1878) at first included it within Pleurotoma purpurea sensu Philippi non Montagu. Thereafter (Monterosato, 1884), he separated to two species and introduced the replacement name Cordieria pupoides noticing an alleged homonymy with “P. rudis Broderip”. Ac- tually, Broderip introduced, in 1834, Placunanomia rudis (a bivalve), the abbreviation P rudis having possibly mislead Monterosato. However, Pleuro- toma rudis Sacchi is preoccupied by P. rudis G.B. Sowerby I, 1 834 (currently accepted as Crassispira rudis ) and by P rudis Philippi, 1836 (currently accepted as Clathromangelia granum (Philippi, 1844): note that Philippi’s work preceeds Scacchi’s one according to Cretella et al., 2005: 115), and the replacement name by Monterosato still holds valid. Regrettably, the type material of Pleurotoma rudis Scacchi is lost (Cretella et al., 2005: 123) and we have established hereby a neotype based on Monterosato ’s material. The original material of Pleurotoma rudis Scacchi has gone lost. We des- ignate, for the sake of stability, a shell from the Monterosato collection, upon which he based his concept of Cordieria pupoides , as the neotype of Pleurotoma rudis Scacchi. Some Authors (Bucquoy et al., 1883: 93) in- cluded, in the synonymy of R. rudis , Pleurotoma reticulata var. brevis Requien, 1848. However, this is a nomen nudum and thus, not available. Nordsieck (1977) used this name ( brevis ) and provided the first valid introduction, but referring to a distinct species. Nordsieck (1968: 176) split R. rudis Scacchi into four subspecies: R. rudis rudis, R. rudis pupoidea [sic!], R. rudis cylindrica and R. rudis intermedia. Descriptions of R. rudis rudis and R. rudis pupoidea [sic! emor pro pupoides ] are quite similar and migth be referred to the same species (R. pupoides). Con- cerning the two other “subspecies”, R. cylindrica (erroneously ascribed to Monterosato, actually introduced by Locard & Caziot, 1899) is a distinct unrelated species; “R. rudis intermedia n. ssp.” had a scanty description and was not figured. Sub- sequently, Nordsieck raised it to species level and provided a description and figure of R. intermedia (Nordsieck, 1977: 56, pi. 18 fig. 140). This is R. laviae, as confirmed by the study of a syntype (SMF, sine numero, with autograph Nordsieck’s label). To increase confusion, Nordsieck (1977: 52) also intro- duced R. ( Cyrtoides ) neapolitana as a replacement name pro Pleurotoma rudis Scacchi, 1836 non Bro- derip, evidently neglecting Monterosato’s introduc- tion: R. neapolitana is thus not available. Material on which Nordsieck based his concept of R. neapol- itana (SMF 340337, 3403379 and 340338) included small size specimens of R. laviae and R. bicolor. Raphitoma cfr. pupoides as figured by Cavallo & Repetto (1992: 147 fig. 401) andR. cfr. pupoides as figured by Cachia et al. (2001: 69 pi. 10 fig. 9) are not referable to the present species. Raphitoma pupoides as figured by Scuderi & Terlizzi (2012: pi. XVIII n. 6) is rather to be referred to R. cordieri sensu Auctores. Raphitoma pupoides can be easily distinguished from R. echinata sensu Auctores by its cyrtoconoid not stepped outline and the shorter siphonal canal. Specimens of R. pupoides with strong sculpture on the last whorls may be confused with R. radula, which is however diagnosed by its more acute spire, the ligther colour without blotches or spots. Mediterranean Raphitomidae, 3: on the Raphitoma pupoides complex, with the description of a new species 107 Figures 1-8. Shells of Raphitoma pupoides (Monterosato, 1884). Fig. 1: Lectotype: Provenza, (MCZR lotto 16492), h: 18.7 mm with label of the lot; Fig. 2: sine locus (MNHN-IM-2000-3240), h: 16.5 mm; Fig. 3: Palinuro, close-up of the sculpture; Fig. 4: Anzio, h: 20 mm; Fig. 5: Anzio, h: 17 mm; Fig. 6: Jesolo (Venezia), h: 20 mm; Fig. 7: Saint-Raphael, Est La Chretienne (France), h: 15.7; Fig. 8: Isola delle Femmine (Palermo), juveniles, h: 9.1 mm. 108 Francesco Pusateri etalii Figure 9. Raphitoma pupoides (Monterosato, 1884), Adriatic, h: 12 mm. Figure 10. Raphitoma radula (Monterosato, 1884), Palermo, coll. Melville-Tomlin, NMW, h: 11.5 mm, with label. Raphitoma alida Pusateri et Giannuzzi-Savelli n. sp. - Figs. 11-15, 25 Examined material. Holotype and 3 paratypes from Palermo (coll. Monterosato, MCZR 16905), with handwritten Monterosato label: “K tomentosal Monts./Palermo”; 2 paratypes, Gulf of Palermo (PUS). Other examined material. Italy. Gulf of Bar- atti, 1 sh (MAR), 1 sh (BOG); Livorno, 1 sh (BOG); Scilla (Reggio Calabria), 3 sh (VAZ); Palermo, 1 sh sub nomine ms. “ perfecta ” (coll. Monterosato, 16905); sine loco probably Palermo, 1 sh, (coll. Monterosato, MCZR 16905); Gulf of Palermo, 2 sh (PUS). “Coste d’Africa”. 1 sh (coll. Monterosato, MCZR 16905). Description of holotype. Shell of medium size for the genus, height 17.1 mm, width 7 mm, fusi- form-pupoid, slender, h/d 2.44 mm. Protoconch paucispiral, only protoconch I of of 1.5 convex whorls, height 540 pm, width 480 pm; sculpture orthogonally cancellate. Teleoconch of 7 convex whorls. Suture not incised, evident. Axial sculpture of 1 6 sligthly opisthocline (somethimes orthocline), elevated and strong ribs, and interspaces twices as broad as the ribs. Spiral sculpture on the last whorl of 6 cordlets, thinner that axial ribs and interspaces twices as broad as the cordlets. Cancellation rectan- gular, with spinulose tubercles at the intersections. Secondary cordlets appearing occasionally and thereafter becoming as strong as the others. Subsutural ramp wide, devoid of evident sculpture. Columella simple, slightly sinuous anteriorly, gently angled posteriorly. Outer lip thickened and crenulated externally, with 9 strong inner denticles, the most posterior smaller, delimiting the wide and deep anal sinus, the most anterior more robust and delimiting the funnel-like siphonal canal. Siphonal fasciole of 7 nodulose cordlets, neatly spaced from the last spiral cordlet. Colour straw yellow, becom- ing gradually orange-brownish in the subsutural area, and with an orange-brown band visible inside the aperture. Comma-shaped white spots on the subsutural ramp, arrow-like white spots inside some cancellation interspaces. Soft parts are unknown. Variability. Paratypes shells: height 12-17 mm (mean 14.4, std 1.66), width 5.5-7 mm (mean 6.36, Mediterranean Raphitomidae, 3: on the Raphitoma pupoides complex, with the description of a new species 109 Figures 11-14. Shells of Raphitoma alida n. sp. Fig. 11: Holotype, Palermo (coll. Monterosato MCZ, lot 16905), h: 17.1 mm; Fig. 12: Paratype A, Palermo (coll. Monterosato MCZR, lot 16905), h: 14.8 mm; Fig. 13: Paratype E, Gulf of Palermo, (PUS n. 405), h: 12.1 mm (sz = subsutural zone; sc = secondary cordlet); Fig. 14: Gulf of Palermo, h: 12.8 mm. Figure 15. Raphitoma alida n.sp., protoconch of the holotype. 110 Francesco Pusateri etalii std 0.57), h/d 2.12-2.36 mm (mean 2.26, std 0.10); axial sculpture of 14-16 ribs; outer lip with 9-10 denticles. Soft parts are unknown. Etymology. From the two granddaughters of the authors (Alice Giannuzzi Savelli and Ida Pusateri), ali[ce]+ida, used as a noun in apposition. Distribution. This new species is known only for the examined material, from Tyrrhenian and Central Mediterranean. Type locality is Palermo. Remarks. Raphitoma alida n. sp. differ from R. pupoides mainly in its paucispiral protoconch (v. multispiral in R. pupoides ). Shells without proto- conch of the new species could be confused with shells of R. pupoides with a non-obsolete sculpture on the last whorl; R. alida n. sp. can be distin- guished by its different background colour (chestnut v. yellowish), 7 nodulose cordlet on the fasciole v. 6 less nodulose in R. pupoides , and the less pupoid and more fusiform outline. Some recent Authors (Nordsieck, 1968, 1977; Piani, 1980) erroneously ascribed to Monterosato a validly published “ Raphitoma tomentosa”. Although the epithet “ tomentosa ” was evidently especially liked by Monterosato, he has never published such binomen. The epithet " tomentosa " was, for mysterious reasons, to be particularly dear and pleasing to Monterosato so that in schedis, gave this name to various entities: - Philbertia tomentosa , lot 16682 = some mixed specimens of R. philberti var. - D. tomentosa, lotto 16901 = 4 specimens of R. horrida. - P tomentosa lotto 16696 = 5 spe- cimens of R. lineolata. - Philbertia tomentosa, Monterosato’s label in coll Coen lot 1912 = 2 spe- cimens of R. pruinosa. Nordsieck (1968: 177) reported Raphitoma phil- berti tomentosa with a useless scanty description (“ kleiner ; gedrungen mit konvexen Umgangen. Schlanker stiel. Hell reh-weiss ”; small, stout, with convex whorls. Slender tail. Light fawn and white) and without any figure. Nordsieck (1977: 58 n. A 149) again reported Raphitoma (Philbertia) to- mentosa ascribing it to Monterosato, 1884, with an apparently good description and a figure (Nord- sieck, 1977: pis. 19 n. 149). However, the four lots labelled under this name in the coll. Nordsieck inckluded the following materials: SMF 341803/1, labelled “ Philbertia tomentosa Mtrs. Egina”, one worn shell, 5.4 mm long, with two holes, protoconch missing, probably R. laviae; SMF 341804/1, labelled “ Philbertia tomentosa Mtrs. Karpathos”, one very worn shell, 3.2 mm long, protoconch missing, probably R. bicolor juv.; SMF 341805/1, labelled “ Philbertia tomentosa Mtrs, Cataldo (Brindisi)”, one very worn shell, 5.9 mm long, protoronch missing, indeterminable. Nordsieck (1977: 58) reported “Palermo, Cataldo”: although there is a beach called San Cataldo near Terrasini (Palermo), it is more likely that the true locality was San Cataldo, not far from Brindisi; SMF 341802/5, labelled as “ Philbertia tomentosa Mtrs., Ibiza”, 5 shells, 2. 5-6. 5 mm long, four too worn to be identified, one referable to R. bicolor juv., with a portion of multispiral protoconch. None of these shells matched the description, the size (7 x 3.2 mm) or the figure provided by Nordsieck, including the described paucispiral protoconch, whilst all but one shells (with traces of multispiral protoconch) lacked the apex. It is worthy of notice that Nordsieck's "descriptions" were not necessarily based (only) on actual speci- mens but frequently included also a compilation from literature. Same holds for his drawings, often compound artwork of actual specimens and figures from the literature. This explains why so rarely spe- cimens can be found which match his figures (our unpublished observations and R. Janssen, SMF, personal communication). Nordsieck included this entity in the subgenus Philbertia, which in his scheme comprised species (R. philberti, R. laviae, R. lineolata, R. atropurpurea, R. densa, etc.) that have nothing to do with the R. pupoides- complex. Parenzan (1970: 212 n. 862) cited R. philberti var. tomentosa Monterosato evidently mutuating it after Nordsieck (1968). This name is anyway unavail- able, having been introduced as a varietal name after 1960 (ICZN, 1999: art. 15.2). Raphitoma radula (Monterosato, 1884) [Cordieria] Figs. 10, 16-23,26 Cordieria radula Monterosato, 1884: 132 Clathurella radula de Monterosato, Locard, 1886: 117 Clathurella radula de Monterosato, Locard, 1891: 67 Clathurella radula de Monterosato, Locard & Caziot, 1899: 250 Clathurella radula var. elongata, Locard & Caziot, 1899: 250 (nomen nudum) Mediterranean Raphitomidae, 3: on the Raphitoma pupoides complex, with the description of a new species 111 Figures 16-22. Shells of Raphitoma radula (Monterosato, 1884). Fig. 16: Lectotype, Palermo, (MCZR), h: 14.8 mm; Fig. 17: particular (sc = secondary cordlet); Fig. 18: Palermo (coll. Monterosato MCZR), Paralectotype A, h: 17 mm; Fig. 19: Isola d’Elba, h: 1 8 mm; Fig. 20; Palermo (coll. Monterosato MCZR), Paralectotype F, h: 6 mm; Fig. 2 1 : Antignano (Livorno), h: 9.9 mm; Fig. 22: Gulf of Palermo, h: 12.7 mm. Figure 23. Raphitoma radula , protoconch of paralectotype F. 112 Francesco Pusateri etalii Figures 24-26. Siphonal fasciole of Raphitoma pupoides (Fig. 24), R. alida (Fig. 25), and R. radula (Fig. 26). Clathurella radula var. fuscescens, Locard & Caziot, 1899: 250 (nomen nudum) Clathurella radula var. lutescens, Locard & Caziot, 1899: 250 (nomen nudum) Clathurella radula var. minor , Locard & Caziot, 1899: 250 (nomen nudum) Clathurella radula var. ventricosa , Locard & Caziot, 1899: 250 (nomen nudum) Cordieria radula Monterosato, Pallary, 1900: 256 Raphitoma reticulata radula Nordsieck, 1968: 175, pi. 29 fig. 94.16 Raphitoma echinata cordieri form d (radula) Monterosato, Nordsieck, 1977: 51 Cordieria radula (Monterosato), Sabelli et al., 1990: 217 Type locality. Palermo. Examined material. Lectotype (here design- ated, 14.8 x 6.4 mm) Monterosato coll (MCZR 16476), with handwritten label by Monterosato “Cordieria/ radula, Monts/Nomencl. p. 132/Palenuo”; and 11 paralectotypes Monterosato coll (MCZR 16476) with handwritten label by Monterosato “C. radula/ Pal!!”. Spain. Alboran, -80 m, 1 sh, (SBR); Cadiz, 1 sh (MNHN). France. St. Henry (Marseille), 4 sh (coll. Locard MNHN); Marseille, 5 sh (coll. Locard MNHN); Toulon, 1 sh (coll. Locard MNHN); St. Raphael, 1 sh (coll. Locard MNHN); Sete, 2 sh (coll. Locard MNHN). Italy. Secca delle Vedove, -120/130 m, 2 sh (PAO); Castiglioncello (Livorno), 1 sh (MAR); Capraia Isl., 1 sh (BOG); Napoli, 2 sh (coll. Monterosato, MCZR, sine numero, sub nomine ms. var. aspera); Puolo (Napoli), 1 sh (DUR); Ischia Isl., 1 sh (TRI); Gulf of Palermo, 10 sh (PUS); Gulf of Palermo, 3 sh (coll. Monterosato, MCZR lot 16492, 3), 2 sh (coll. Monterosato, MCZR lot 17342); Porto di Palermo, 2 sh (coll. Monterosato, MCZR lot 16476); Palermo, 3 sh (coll. Melville- Tomlin, NMW); Mondello (Palermo), 1 sh (coll. Monterosato sine numero, sub nomine “ purpurea albino ”); Sciacca, 1 sh (coll. Monterosato, MCZR lot 16492); Catania, (coll. Monterosato ex Aradas, MCZR, lot 16476, 2 sh). Algeria. Sine loco, 2 sh (coll. Monterosato, MCZR lot 16492); Orano, 1 sh (coll. Pallary MNHN). Croatia. Between Pula and Ligthouse of Porer, 1 sh, legit W. Koers (SMNH lot 70484). Mediterranean Raphitomidae, 3: on the Raphitoma pupoides complex, with the description of a new species 113 Description. In squared parentheses data of the lectotype. Shell of medium size for the genus, height 9-19 mm [14.8] (mean 13.81, std 2.90), width 4-8 mm [6.4] (mean 5.90, std 1.10), fusi- form-pupoid, slender, h/d 2. 2-2. 5 [2.31] (mean 2.32, std 0.09). Protoconch multispiral of 2.7 con- vex whorls, height 580 pm, width 440 pm; proto- conch I of 1.1 whorls, width 210 pm, with irregularly placed small tubercles and orthogonally cancellate sculpture; protoconch II of 1.6 whorls, with a diagonally cancellate sculpture. Teleoconch of 7-8 [7] convex whorls. Suture not impressed. Axial sculpture of 12-17 [16] sligthly opisthocline, elevate, strong ribs, and interspaces as broad as the ribs (or sligthly broader). Growth lines visible between the ribs on the last whorl. Spiral sculpture on the last whorl of 5-6 [5] cordlets above the aperture, thinner than axial ribs, with interspaces three times as broad as the cordlets, and a secondary cordlet bordering the subsutural ramp. Cancellation squared. Secondary cordlets appearing occasionally and thereafter becoming as strong as the others. Subsutural ramp narrow, devoid of evident sculp- ture. Columella simple, slightly sinuous anteriorly, gently angled posteriorly. Outer lip thickened and crenulated externally, with 8-9 [9] (rarely up to 11) strong inner denticles, the most posterior smaller, delimiting the wide and deep anal sinus, the most anterior more robust and delimiting the funnel-like siphonal canal. Siphonal fasciole of 7-8 [7] nodu- lose cordlets, neatly spaced from the last spiral cordlet. Colour from uniformly whitish to very ligth chestnut brown, with darker subsutural ramp and darker band on the lower part of the last whorl. Violet hue on the background in particularly fresh specimens. Comma-shaped white spots on the sub- sutural ramp, arrow-like white spots inside some cancellation interspaces. Soft parts are unknown. Distribution. Provence, Western Mediter- ranean and Tyrrhenian. A single record from neigh- bouring Atlantic (Cadiz). Remarks. Raphitoma radula could be confused with shells of R. pupoides with non-obsolescent sculpture, but it is easily diagnosed by its homo- geneous ligth coloration with violet hue. It could me mixed with very ligth or albinistic shells of R. echinata (of similar size) from which it differs in the less elevate spirals, the shorter and more roun- ded aperture and the violet hue in fresh specimens. Monterosato (1884: 132) introduced Cordieria radula for the erroneously identified P. pur- pureum sensu Philippi (non Mtg.), referring to the examen (ex typo) of a specimen provided by Philippi himself to Sylvanus Hanley. According to Clare Brown (Leeds Museum Discovery Centre) “Hanley’s collection came to us [LMG- NS] in the 1950s after being broken up and many parts sold on. Sadly, it seems as if the Philippi P. purpurea didn’t make it to Leeds”. However, there is little doubt that the type material of Cordieria radula Monterosato consists of the type series at MCZR. ACKNOWLEDGEMENTS Roberto Ardovini (Rome, Italy), Cesare Bogi (Livorno, Italy), Philippe Bouchet (MNHN), Clare Brown (LMG-NS), Sergio Duraccio (Napoli, Italy), Jennifer Gallichan (NMW), Alfio Germana (Trecastagni, Catania, Italy), Sandro Gori (Livorno, Italy), Virginie Heros (MNHN), Andre Hoarau (Frejus, France), Piera Iacovelli (MRSNT), Ronald Janssen (SFM), Gabriele Macri (Scorrano, Lecce, Italy), Alessandro Margelli (Livorno, Italy), Paolo Mariottini (Rome, Italy), Henk Mienis (HUJ), Attilio Pagli (Lari, Pisa, Italy), Paolo Paolini (Livorno, Italy), Jakov Prkic (Split, Croatia), Paolo Russo (Venezia, Italy), Carlo Sbrana (Livorno, Italy), Gabriele Sercia (Palermo, Italy), Carlo Smriglio (Rome, Italy), Peter Sossi (Trieste, Italy), Gianni Spada (Vagrigneuse, France), Ennio Squizzato (Loreggia, Padova, Italy), Morena Tisselli (S. Zaccaria, Ravenna, Italy), Lionello Tringali (Rome, Italy), Evi Vardala-Theodoru (Athens, Greece), Angelo Vazzana (Reggio Ca- labria, Italy), Anders Waren (SMNH), provided ma- terial and informations. The staff at the Museo Civico di Zoologia di Roma (MCZR), and particu- larly Director Claudio Manicastri and Curator Massimo Appolloni, continuously supported our researches. Gianni Repetto (Alba, Italy) provided precious bibliographic material. SEM photographs were done at the “LIME” (Interdepartmental Laboratory of Electron Microscopy) by Andrea Di Giulio (Dept, of Biology, “Roma Tre” University, Rome). Marco Oliverio (Rome, Italy) for the crit- ical review of the manuscript and for helpful sug- gestions. 114 Francesco Pusateri etalii REFERENCES Aartsen J. J. van, Menkhorst H.P.M.G. & Gittenberger E., 1984. The marine Mollusca of the Bay of Algeciras, Spain, with general notes on Mitrella, Marginellidae and Turridae. Basteria, suppl. 2: 1-135. Aradas A. & Benoit L., 1872-1876. 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Fischer, Cassel, 512 pp. Biodiversity Journal, 2016, 7 (1): 117-198 Monograph On the origin of allopatric primate species Marc G.M. van Roosmalen* &Tomas van Roosmalen ‘MVRS - Marc van Roosmalen Stichting, Leiden, The Nether lands Corresponding author, e-m ail: marc.mvrs@gmail.com ABSTRACT Here we present a theory on the origin of allopatric primate species that follows - at least in Neotropical primates - the irreversible trend to albinotic skin and coat color, called “meta- chromic bleaching”. It explains why primates constitute such an exceptionally diverse, species-rich, and colorful Order in the Class Mammalia. The theory is in tune with the principle of evolutionary change in tegumentary colors called “m etachrom ism ”, a hypothesis propounded by the late Philip Hershkovitz. Metachromism holds the evolutionary change in hair, skin, and eye melanins following an orderly and irreversible sequence that ends in loss of pigment becoming albinotic, cream to silvery or white. In about all extant sociable Neo- tropical monkeys we identified an irreversible trend according to which metachromic varieties depart from the saturated eumelanin (agouti, black or blackish brown) archetypic form and then speciate into allopatric taxa following the trend to albinotic skin and coat color. Speci- ation goes either along the eumelanin pathway (from gray to silvery to cream to white), or the pheomelanin pathway (from red to orange to yellow to white), or a combination of the two. The theory represents a new and original evolutionary concept that seems to act indef- initely in a non-adaptive way in the population dynamics of male-hierarchic societies of all sociable primates that defend a common territory. We have successfully tested the theory in all 19 extant Neotropical monkey genera. Our theory suggests the trend to allopatry among metachromic varieties in a social group or population to be the principal behavioral factor that empowers metachromic processes in sociable N eo tropical monkeys. It may well represent the principal mechanism behind speciation, radiation, niche separation, and phy logeography in all sociable primates that hold male-defended territories. We urge field biologists who study primate distributions, demography and phy logeography in the Old World to take our theory to the test in the equally colorful Catarrhini. KEYWORDS Neotropical primates; phylogeography ; metachromic bleaching; speciation; radiation. Received 20.12.2015; accepted 09.02.2016; printed 30.03.2016 INTRODUCTION We could ask ourselves (Darwin, 1 859): “ Why primates constitute by far the most diverse, species- rich and colorful Order in the Class Mammalia? Do primate diversity, metachromism and meta- chromic processes relate directly to sexual selection ? Or, rather to its generally complex, hierarchically organized social structure and male territoriality? If not sexual selection, what could be the principal factor(s) in primate social behavior to be held responsible for metachromic processes, speciation, radiation, niche separation, and phylogeography?” Inspired by A lfred Russel Wallace whose concept of the “Origin of Species” was laid down in a paper 118 Marc G.M. van Roosmalen & Tomas van Roosmalen he sent for review to Charles Darwin, here we introduce a new and original theory about species evolution taking place in particular in sociable territorial prim ates. Our theory “ On the tendency of metachromic varieties in sociable primates to depart indefinitely from the agouti archetype and evolve in advanced eumelanin, pheomelanin to albinotic bleached allopatric taxa” is equally rooted in life-long fieldwork on socio-ecology of all Neo- tropical monkey genera, both in captivity and in the wild. It closely follows the principle of evolutionary change in tegumentary colors called “meta- chromism”, a hypothesis propounded by Philip Hershkovitz (1968; 1977).Metachromism holds the evolutionary change in hair, skin, and eye melanins following an orderly and irreversible sequence that ends in loss of pigment through which a taxon of a given genus or phylogenetic clade eventually be- comes albinotic, cream to white. Individual hair color or the entire coat changes from agouti (char- acterized by alternating blackish-brown and reddish bands on the terminal half of the hair) to uniformly blackish -brow n , and thereafter to gray, and even- tually to white or colorless, called the eumelanin pathway; or, it changes from agouti to uniformly reddish to orange to yellow to cream, and even- tually white, called the pheomelanin pathway. The process itself is called saturation, which means the change from the primitive agouti pattern of the hair, or part of the pelage, or the entire coat, to a satur- ated eumelanin (blackish) or saturated pheomelanin (reddish) coloration. The dilution, or gradual reduc- tion in the amount of pigment deposited in the growing hair, and disappearance of pigmentary colors is called bleaching (Fig. 1). In the color of the skin and iris of the eye, it follows the eumelanin pathway (brown to drab, to gray, or blue), and then it is termed depigmentation. Metachromism applies to all mammalian species. It is thought to also occur in bird feathers. Our theory suggests that among social groups or populations of advanced intelligent, socially organized, male-territorial mammals, in particular primates, phenotypical varieties (mutants) that show slightly bleached eumelanin or pheomelanin colored skin or coat characteristics do arise indef- initely. Their melanocytes (skin cells that produce the black pigment melanin) are smaller, and for that or any other reason produce less melanin. In gen- eral, the tendency of these metachromic varieties is Figure 1. Bleaching from saturated eumelanin and saturated pheomelanin fie Ids to white or colorless. Gradual reduction in the amount of pigment deposited in the growing hair re- sults in apparent change from blackish through brown, drab gray to white or colorless in the eumelanin pathway, and from reddish through orange, yellow, cream to white in the pheomelanin pathway. Switching from the eumelanin to the pheomelanin pathway occurs in saturation but not in blea- ching (modified from Hershkovitz, 1977). neotonic, taking place locally (e.g., naked muzzle, bald head, euchromic blaze/forehead or part of the coat, depilation of skin) or all over the body. Social structure in most primate societies, in particular those of the more advanced monkeys and apes, is hierarchically organized, whereas male over female dominance is the rule, with very few exceptions (e.g., spider monkeys in the Neotropics and pygmy chimpanzees in Central Africa adopted a matri- archal social system, in which males patrol and defend a common territory, and females are allowed to transfer to neighboring social groups). Social selection is the recognition of and preference for the parental (or foster parental) phenotype in societal grouping and mating. Social selection for color or color pattern through assortative mating tends to stabilize within a chromatic range recognized and accepted by free-ranging but chrom otypically imprinted members of the social group. Slightly depilated or somewhat eumelanin or pheomelanin bleached individuals deviant from the socially selected skin or hair color pattern, in particular when it is detected in adolescent to subadult males, may be discriminated against by high-ranking (alpha)-m ales. For that reason alone they can be pushed into the periphery of the group. Depending on the primate taxon or genus, such individual young males may also be expelled from the parental On the origin of allopatric primate species 119 group, and then become social ‘outcasts’. Peri- pheral or outcast males do suffer on a daily base from less and shorter access to the group’s prefer- ential, comparatively more nutritious food sources. They may join one another for reasons of social comfort and during ranging or foraging they tend to hang out together at the periphery of the group. Eventually, they may decide to leave the pack as all-male parties and roam around in much larger areas than just the home range or territory of the parental group. They then may attract young fe- males from neighboring social groups. Together, they may seek some hitherto overlooked, ‘empty’ or little-used living space in an attempt to settle down and start their own family or social group. In case the taxon or genus it belongs to shows ter- ritorial behavior - which is the case in almost all Neotropical monkeys - these emigrants will be sub- sequently pushed out from neighboring territories as well. Consequently, they will die from starvation, parasite load and/or diseases forthcoming the dietary constraints they are suffering from. Or, as a mat ter of luck, in the end they may find some living space that is not (yet) occupied, most likely at considerable distance from the taxon’s core distri- bution. Sometimes, such emigrant parties can be forced to survive in peripheral habitat that has to be considered marginal for that species to occur in. In extremely rare cases, such parties might even manage to circumvent a certain geographical barrier and beyond it find for the species appropriate hab- itat, where their specific ecological niche is not occupied, as such involving a range extension. In case that habitat is already occupied by a closely related species, a battle for life will take place and the best fitted taxon will drive the other to extinc- tion, the red-handed tarn arin Saguinus lTlidas that is replacing the bare-face tam arin S. bicolov. Only over geological spans of time, for example after a vicariance has taken place, suitable habitat may open up where the taxon’s niche is not occupied by another primate. One may imagine that along these paths sm all reproductively isolated founder-colon- ies that contain somewhat bleached and/or depilated individuals may establish them selves there. For the sake of survival alone they would unselectively interbreed or hybridize. Inbreeding then may relax stabilizing forces and stimulate or accelerate meta- chromic and other degenerative (= non-adaptive) processes. The more metachromic advanced each successively isolated breeding colony is, or the farther it has moved from the center of that taxon’s dispersion, the nearer it will come to the end of its metachromic evolution. And, the narrower will be its range of chromic fitness (e.g., prey and predator camouflage). This degenerative process, though, may be counterbalanced if under strong natural selection newly diverged forms, that evolve in a, for the original species marginal or new habitat, niche or landscape, at the same time selectively become better fitted, more cooperative, more inventive, or smarter in the adaptation process. This may happen every time founder-colonies successfully travel across existing geographical barriers, such as rivers, watersheds, mountain ranges, or open areas with arid scrub vegetation. Completion of the processes of metachromic bleaching, depigmentation, or de- pilation, whether taking place single or combined, eventually will result in extinction of the race or species. Unless the founder-colony or population in time does find and manage to occupy hitherto empty, but suitable habitat. Or: if it adapts to a dif- ferent ecological niche, where skin and coat color do not have survival value by lack of competition from closely related species. Dead-end, isolated, peripheral, or new habitats may be occupied by metachromic dead-end populations, such as has happened over and over in the Neotropics in ad- vanced albinotic callitrichids, uakaris, sakis, titis, capuchins, howlers, spider, woolly, and woolly spider m onkey s. RESULTS In non-territorial, peaceable Dwarf Marmosets Callibella humilis neotony and euchromism are clearly demonstrated as infants are overall much lighter colored than adults, showing a tendency to albinotic. Their overall pelage is light brown, their tail alternately light and dark-brown banded, and their face flesh-colored with a circumference of long, bright white hairs. From three months on, they pass through a complete metachromic metamorph- osis. Their overall coat turns into saturated eu- melanin, the muzzle of their faces into pinkish, and their semicrescent ocular rings or eyebrows into white (Fig. 2). This natural process may be related to slightly smaller melanocytes (skin cells that produce the black pigment melanin) producing overall less melanin. 120 Marc G.M. van Roosmalen & Tomas van Roosmalen Callibella stands at the base of the phylogenetic tree from which all extant A m azonian marmosets, Cebuella and Mico, have derived (Van Roosmalen & Van Roosmalen, 2003). It finds itself at the verge of extinction, for it occupies the niche of exudate gouging - that is feeding on resins ouzing out of little holes they themselves have gnawed in the bark of certain gum trees and climbers. That niche is filled in by the advanced, larger, highly territorial Amazonian marmoset genus Mico (its distributions are shown in figures 5-7). We believe that these aggressive, over twice as big callitrich id monkeys have displaced the non-territorial dwarf marmoset and taken over its specific feeding niche all over its former, much larger range - the entire interfluve delineated by the Rios Madeira, Amazonas and Tapajos. The genus Callibella is thought to have evolved there in the late-Pliocene to early-Pleisto- cene landscape that was dominated by lacustrine seasonally inundated clear-water igapo wetlands. Being peaceable monkeys that like their neighbor’s company instead of attacking or trying to kill them apparently has not been an evolutionary success among primates (Van Roosmalen, 2013a, b; 2015). Contrastingly, the pygmy marmoset Cebuella that derived from prototypic Callibella nowadays occupies the entire western Amazon Basin. We believe it was so successful because Cebuella , being allopatric with Callibella and Mico, could occupy the ecological feeding niche of exudate gouger west of the Rio Madeira. There, it did not have to face competition from other callitrichids over exudate food sources. Indeed, Amazonian tamarins (genus Saguinus) that range west of the Rio Madeira (Figs. 7-13) lack the elongated tusked mandibular second incisors needed for tapping sap Figure 2. Ontogeny in Black-crowned D w arf M arm o sets Callibella hwnilis V an Roosmalen & Van Roosmalen (2003) based on photos of captive and wild individuals (Van Roosmalen et al., 1998). On the origin of allopatric primate species 121 from tree barks. As such, tamarins do not directly compete with pygmy marmosets over gum. Instead, tamarins of different taxa are reported to parasitize on pygmy marmosets by licking the resins from tap holes made by the latter. Cebuella pygmaea being overall agouti colored is clearly the most archetypic among the two extant taxa of pygmy marmoset. Distributed north of the Upper Amazon River(Rio Solimoes/Rio Maranon) and specialized in exudate gouging, the species (or its precursor) seems to have adapted to seasonally white-water inundated floodplain forest (varzea) habitat. Somewhat pheomelanin bleached colon- izers of ancestral C. pygmaea (having an orange colored tail and breast, progressively bleached yel- low to white belly, yellow-white mustache, naked pink-colored muzzle and circumocular rings) fol- lowing the trend to allopatry once must have man- aged to traverse the Amazon River proper, on floa- ting varzea islands and/or passively through ri- verbend cut-offs (oxbow lakes). By lack of competitors the nearest to albinotic taxon Cebuella tliveiventris - the form that derived from allopatric archetypic C. pygmaea - could then have extended its range from the Rios Javari and Jurua east as far as the Rio Madeira and south of the Amazon River as far as the Bolivian Amazon (Fig. 3). There, it secondarily adapted to never inundating terra firm e high forest. Nowadays, it is found there, especially at edges of treefall clearings and in secondary growth. As C. tliveiventris is fully allopatric with C. pygmaea and, moreover, shows completely dif- ferent habitat preferences, we here propose to at- tribute both taxa full-species status naming them C. pygmaea and C. niveiventris. During our system- atic surveys of primate distributions and diversity Figure 3. Present-day distributions are here depicted for Black-crowned Dwarf Marmosets genus Ccillibellci and Pygmy M arm osets genus Cebuella, representing the sm allest m onkey s in the w orld .The current distribution of the m onotypic genus Callibella perhaps has to be considered the sm allest of any prim ate in the N eotropics. 122 Marc G.M. van Roosmalen & Tomas van Roosmalen carried out in the matrix terra firme hinterland stretching out behind the floodplain of white-water rivers (i.e., the Rios Javari, Jurua, Purus and Madeira), we were not able to detect any pheno- typical difference between individuals sighted at any point along these far-apart rivers. It may indicate that in highly territorial monkeys like pygmy marmosets that occupy large distributions delineated by some of the largest tributaries of the Amazon River, phenotypical characters of skin and pelage coloration and/or local hair growth or de- pilation seem to have stabilized. In other words, we believe that within a given monkey’s distribution something like a gradient of slightly differing phenotypes or color morphs, or geographic races, in reality does not exist. These and other observa- tions from the larger field have led us to attribute full-species status to monkey taxa such as C. pygmaea and C. niveiventris that we ourselves have confirmed to be phenoty pically stable throughout their (sometimes very large) range. Here, we would like to propose a new species concept: ecospecies. This species concept is further corroborated by the here introduced evolution theory that aims to explain the origin of allopatric primate species. We define ecospecies as follows: "An ecospecies is a genetically isolated population or group of populations of a kind that does not undergo any gene flow from other populations of one or more closely related kinds, and that demon- strates a stabilized, well-defined phenotype over its entire range, in which it occupies and defends a specific ecological feeding niche against any out- side Competitor'’'. This definition of a primate species avoids the confusing, rather arbitrary dis- tinction between species and subspecies (or race), for it adds sociobiological factors to geographical, geom orphological and phy tosociological ones that act on the evolutionary process of primate speci- ation and radiation. Following this concept, for in- stance, an enclave population of Callibella humilis that we found living year-round in the seasonally inundated floodplain forest (igapo) along both banks of the Rio Atininga - genetically isolated from the main population occuring in terra firme forest at least one-hundred km to the north - should be given its own species name and treated as such. Or, in case the ranges of two saddle-back tamarins of the SaguinUS fuscicollis Clad e, hitherto taxonom- ically treated as subspecies, are separated by a con- tact zone, where territorial behavior effectively im- pedes gene flow through hybridizing, both popula- tions should be attributed full-species status. The c allitric h id s Goeldi’s Monkey Callimico and Black-crowned D w arf M arm o set Callibella do represent the only monospecific (= monotypic) primate genera in the Neotropics. Callimico lives in the upper Amazon Basin region of Bolivia, Brazil, Peru, Colombia, and Ecuador (Fig. 4). Goeldi’s monkey coat coloration is saturated eu- melanin, blackish or blackish-brown. It forages in dense scrubby undergrowth of low mixed forests with discontinuous canopies and in so-called ‘tabocais’ (low forest dominated by bamboo) at levels of less than five meters. Social groups consist- ing of monogamous pairs with single offspring count on average six individuals. Groups live in patches of suitable habitat, often separated by miles of unsuitable vegetation. Goeldi’s monkeys are vertical dingers and leapers able to leap horizontal distances of up to four meters between branches. As they are peaceable monkeys not showing any form of territoriality, Goeldi’s monkeys often associate in mixed species groups with different species of tamarin SaguinUs (Mittermeier et al., 2013). The fact that this primitive little monkey, just like the dwarf marmoset Callibella, remained archetypic in its blackish agouti coat coloration, is peaceable, is not showing any territorial behavior towards its neighbors, is occupying a unique feeding niche (foraging on the ground for fungi and invertebrates, and for fruits at low levels of a discontinuous canopy), and over geological time-span did not diverge into more than one taxon, strongly supports our doctrine that attributes speciation and radiation in m ale -territorial Neotropical primates primarily to the trend to allopatry as expressed in meta- chromic bleaching. As shown in the schematic distribution map of all known Amazonian marmosets (Fig. 5), each in- terfluve in the area delineated by the most effective riverine barriers - Rio Amazonas in the north, Rio Madeira in the west, Rio Guapore in the south, and Rios Tapajos-Juruena and Xingu in the east- is inhabited by a different taxon of MicO, which species phylogeographically and phylogenetically radiated away from an ancestral, archetypic agouti- colored form much resembling the extant species M. melanuruS (Van Roosmalen et al., 2000) from the upper Rio Aripuana basin - the taxon with the On the origin of allopatric primate species 123 southernmost distribution of all Amazonian mar- mosets to be placed at the base of MicO’s phylo- genetic tree. Four monophyletic cladistic Groups or Clades are distinguished: the B are-ear M. OTgentatUS C lade, the (Tufted-ear or) Tassel-ear M. hwTieralifer Clade, the W hite-m an tie (w hite-hip) M. melcMUruS Clade, and the Orange-leg M. tnarcai Clade (Figs. 6, 7). Within each Clade, the evolutionary pathway towards advanced metachromic bleached (and ultimately albinotic) taxa can be plausibly retraced. Albinotic forms in dead-end distributions may eventually go extinct (i.e., M. chrysoleuCOS in the M. humeralifer Clade; the new Mico species that occurs between the Rios Teles-Pires and Ronuro, M. leucippe, and M. argentatus in the M. argentatus Clade; M. acariensis and M. saterei in the M. melanurus Clade; and M. manicorensis in the M. marcai Clade). In territorial sociable primates the principle of metachromic bleaching that seems to fuel the trend to allopatry is an irreversible, seem- ingly non-adaptive evolutionary pattern. The meta- chromic pathway followed within the M. argentatus Clade is a predominantly pheomelanin one, with first the nearest to archetypical, dark orange-colored taxon M. emiliae from the Rio Iriri. From M. eifliliae diverged in southward direction the moderately bleached new species that we identified to occur between the Rios Ronuro and Teles-Pires, and northward the advanced albinotic taxa M. leucippe (all white with a pink face) and M. argentatus (all white with a black tail). The latter occupy dead-end distributions, as they are pressed at their northern limit against the u ntraversable Rios Tapajos and Amazonas, respectively. Within the tufted-ear or tassel-ear Clade of Mico the metachromic pattern followed the eumelanin pathway, from the darkest agouti-colored taxon M. mauesi going straight into Figure 4. Present-day distribution of monotypic Goeldi’s Monkey CcillitYlicO goeldii. 124 Marc G.M. van Roosmalen & Tomas van Roosmalen the overall whitish and gray M. hwnercilifer in the Clade’s northernmost dead-end distribution (delin- eated by the Amazon and Tapajos Rivers). Along the pheomelanin pathway it diverged into the overall orange and white colored golden-white tassel-ear marmoset chrysoleucos, the species that occupies the westernmost dead-end distribution delineated by the Rios Madeira and Aripuana. W ithin the w hite-m an tie (w hite-hip) C lade of Mico the pathway followed goes from the nearest ar- chetypic agouti-colored taxon M. melciYllirus in northern direction to the advanced pheomelanin bleached half-way albinotic taxa M. intennediliS , M. acariensis , and M. saterei. And within the fourth Group of Mico, the orange-leg M. MClYCCli Clade, the pathway followed in northern direction starts from the metachromic nearest to archetypic taxon M. ITiarCCli diverging into the advanced euchromic to almost albinotic taxon M. YYlCOflicOYCYlsis , and in western direction proto -lTlCirCCli evolved into the slightly but progressively bleached taxa M. Yligri- CepS and M. VOYldoni, all three occupying dead-end distributions delineated by the untraversable Rio Madeira (after the Amazon River proper the second strongest river barrier in the entire Amazon Basin). As all interfluves occupied by a different Mico species show dead-end distributions delineated by untraversable rivers at their northern and western limits, each species represents a different stage along the eumelanin or pheomelanin pathway that is frozen in time, but at the end of its metachromic evolution it invariably will turn into albinotic (Figs. 6, 7). Once arrived there, such primate taxa will in- evitably go extinct, unless a founder-colony man- ages in time to cross the geographic (riverine) barrier by means of a river bend cut-off, by hopping on varzea forested floating islands, or by circum- venting a geographical barrier. According to the doctrine, the evolutionary rate of metachromism is primarily controlled for by the trend to allopatry, and secondarily by environmental and genetic factors which may accelerate, retard, or terminate metachromic processes, or hold them in dynamic equilibrium, but cannot alter, reverse, or deflect them from their course. Hypothetically, growth and spread of a founder-colony of a certain Amazonian marmoset across a certain interfluve delineated by rivers, entails social selection. Effective selection stabilizes the mean chromotype of the colony at a color tone or grade inbetween that of the founders and that of the albinotic ones towards which all monkeys tend. Amazonian m arm osets (genus Mico ) represent an advanced stock of callitrichids that evolved as late as the Pleistocene, south of the Amazon River and east of the Rio Madeira, from an ancestral stock of the Ccillithrix ouistitis occur- ring in Central and SE Brazil (Van Roosmalen & Van Roosmalen, 2003). About 1.5 MYA, a major vicariance took place - the break-through by the proto-Madeira River of the continental watershed running across the Chapada dos Pareds in Rondo- Tlia (Grabert, 1991). Thereafter, the entire area south of the Amazon and east of the Madeira drastically reversed its drainage pattern. Former rivers that since the beginning of the Pliocene had been drain- ing the extensive clear-water wetlands in north- south direction, dried up. New rivers (mostly of the black-water type) arose and began to drain the area in opposite direction, from south to north. Most of these rivers emptied out in the Rio Madeira, some directly in the Amazon River. Founder-colonies at different phenotypic stages of metachromic bleach- ing that derived from archetypic M. melcinurus - pushed by the trend to allopatry - subsequently invaded and inhabited the newly formed interfluvial terra firm e ‘islands’ that new rivers had been creating. These newly available lands offered them their preferred habitat of terra firme rainforest, in which they filled the niche of exudate gouging, which niche east of the proto-Madeira River was hitherto exclusively occupied by the much smaller and peaceable, non-territorial dwarf marmoset Callibella huinilis (Van Roosmalen & Van Roos- m alen, 2003). Ever since, Ccdlibcllci hwflilis seem s to have lost the battle against the aggressively expanding Mico newcomers. Our assumption is that the dwarf marmoset has been locally driven to extinction almost all over its former range since the genus evolved in the late Pliocene. Presently, the black-crowned dwarf marmoset hangs on along the westbank of the lower Rio Aripuana. As a com- mensal, it takes refuge on the terras pretas (human- made black-earth farmland) from the deadly attacks of the local A m azonian m arm oset Mico MClTlicoreYl- sis (Van Roosmalen et al., 2000; Van Roosmalen & Van Roosmalen, 2003). This example may well demonstrate that a specific ecological niche such as that of specialized gougers and feeders of gum (exudates) in a certain habitat (e.g., primary rain forest) can only and exclusively be occupied by a On the origin of allopatric primate species 125 Amazonas hvmtrrahtrr f nov . Amazonian Marmosets Cebuella Mico Callibella I ^ Figure 5. Schematic distributions as delineated by (for Amazonian Marmosets) un traversable rivers drawn for all known Amazonian Marmosets belonging to the genera Callibella, Cebuella, and Mico. 126 Marc G.M. van Roosmalen & Tomas van Roosmalen ALL PYGMY, DWARF & AMAZONIAN MARMOSETS Cebu elf a (pygmaea) pygmaea and C. (p.) nivetvomris ? 1 %Callibeila humilrs Tassel-ear humeralifer Clade Mico humeralifer t imauesi chrysoleucos Bare-ear argentaws Clade Mico sp, nov, Rio Ronuro emiliae £ argentaws leucippe hite-mantle melanurus Clade Mico melanurus ^ intermedia s m acariensis saierei W Orange-leg marcaf Clade ^ Mico rondoni Qmarcai mantcorensis Ba ^ _ .* "v** 1 4ft Figure 6. Geograph ical distribution s delineated by rivers drawn in one map for all known Amazonian Marmosets that belong to the marmoset genera CcillibellcL, Cebliellci , and Mico. single taxon that defends it, in this case even beyond generic bounds. Figures 5-7 demonstrate that about all inter- fluves occupied by a single species of Mico show dead-end distributions delineated by for rainforest habitat-specialists untraversable rivers at their northern and western limits.At their southern limits, all distributions invariably show a mostly narrow open-end, where a contact zone between two adjacent distributions must exist. Hybridization between Amazonian marmosets, though, has never been seen orreported in the wild. This may well be attributed to strong social and sexual selection. Indeed, all Amazonian marmosets of the genus Mico developed hypertrophied external genitalia in each gender that are physically greatly differing among related taxa (Van Roosmalen et al., 2000) (Figs. 8-11). We ourselves have kept, raised and bred with a number of Amazonian marmosets, both in free- ranging and captive conditions. Expressive and often violent territorial behavior of all members of a social group, aside of species-specific sexual display of external genitalia, pheromones and scent- marking of one another’s coat, has always preven- ted our marmosets from hybridizing (interspecific cross-breeding). For instance, we kept breeding social groups of all three taxa of the tassel-ear M. humeralifer Clade (i.e ., M. humeralifer , M. mauesi, and M. chrysoleUCOS). To avoid one group from wiping out the other, we had to keep different species in separate cages, whereas we let only one group of M. chrySOleUCOS free-ranging in the forest that surrounded the compound. Even so, adults were still seen trying to grab and bite one another through the fine-mess wire. From our unique exper- On the origin of allopatric primate species 127 Cehttella pygi nova argentatus Clade Tassel-ear humeralifet Clade hello hit mil is White-mantle Ml co melon it rtts Clade Orange-leg Mi co mttreai Clade 12tm Figure 7. Radiation and metachromic diversification following eumelanin and pheomelanin pathways of metachromic bleaching depicted for all recognized phylogenetic C lades of Amazonian ( MicO ), D w arf ( CciUibellci) , and Pygmy Marmosets ( Cebuella) depicted to scale. 128 Marc G.M. van Roosmalen & Tomas van Roosmalen Figures 8-11. In all taxa of Mico, both m ales and fem ales evolved hypertrophied . species -specific , in anatomical respect very differently shaped external genitalia. Fig. 8: male M. manicOTensis sexually displaying; Fig. 9: exposed pudenda in adult female M. acarietisis ; Fig. 10: pudenda with 2 cm long vaginal lips in M. SCLterei, Fig. 11: pudenda in M. CLCClriensis. This feature supports our view that all taxa of Mico should be considered different species and not just metachromic color morphs. ience having kept all kinds of marmosets (until today, not a single zoo in the world has any Mico on exhibit) and other callitrichids, both in captivity and free-ranging in a tropical rainforest environ- ment, we believe that where adjacent distributions of two species of Mico are not defined by an un tra- versable river, a sharp-lined contact zone must exist, where cross- breeding never takes place. This assumption concurs with the principle of meta- chromic bleaching being irreversible. In theory, only through cross-breeding with a darker, overall more saturated eumelanin taxon the metachromic pathway to albinotic could be reversed, something, however, that will never happen in the wild. As all Mico do display strong interspecific territorial behavior - each group defending its living space by means of (often ritualized) territorial boundary conflicts - within a given contact zone cross-breeding will not take place between neigh- boring groups of different but related ecospecies, as distance is maintained by regularly performed boundary conflicts. This way, any gene flow between phenotypically different populations is impeded. In phylogeographic terms, the farther radiated away from the origin of a Clade’s disper- sion - that of the nearest to archetypic species within a monophyletic Clade - the more progressively bleached the species will become. Partly or fully albinotic taxa, therefore, often occur in or near the Clade’s dead-end distributions. In figures 12-15, we have visualized the phylo- geographic distributions, radiation, and supposed pathways of metachromic bleaching of all known Tamarin Monkeys genus Sciguinus. We have di- On the origin of allopatric primate species 129 vided them up in the following monophyletic Groups or Clades: the Saddle-back Tamarins of the S. fuscicollis Clade (Fig. 13); the Black-mantle White- mouth Tamarins of the S. nigricollis Clade in one map combined with the Mustached Tamarins of the S. my StttX Clade, the Red-chested Mustached S. labiatUS Clade, and the Emperor Mustached S. imperator Clade (Fig. 14); and the Bare-face Tamar- ins of the S. midas, S. bicolor and S. geoffroyi Clades (Fig. 15). To complete the c a llitric hid picture, we have visualized the distributions of the Fion Tamarins genus LeOYltopitheCUS , and the True Marmosets or Ouistitis genus Callithrix, from SE B razil (Fig. 16). In geological history, speciation and radiation within the Saddle-back Tamarins of the S. juscicol- lis Clade (Figure 13) went along two pheomelanin pathways of metachromic bleaching: one sub-Clade radiated south of the Amazon River from east to west, from the most saturated eumelanin, nearest to archetypic taxon S. MUra (green distribution) to the completely albinotic all-white taxon S. melanoleii- CUS (blue distribution) via the taxa S. avilapiresi, S. fuscicollis, and S. CTUzlimai. The bleaching process took first place in the head parts - muzzle and blaze - and, after having traversed the Rio Jurua back to its right bank, the metachromic bleaching process completed from the overall orange-colored taxon S. cruzlimai into the fully albinotic taxon S. melanoleucus. A nother radiation took place from S. mura directly into S. Wcddclli, and, after having traversed the Rio Purus, into the overall light-brown colored taxon S. primitivUS - both with a fully al- binotic blaze and muzzle/mouth. A second sub- Clade of saddle-back tamarins radiated from the Peruvian Amazon in eastern direction, from the saturated eumelanin nearest to archetypic taxon S. leUCOgenys (light blue distribution) into the slightly ALL PYGMY. DWARF * AMAZONIAN MARMOSETS Cpjrfpn#**) jTppnM* *nd C /* WC**+H**+mm ^2 ■*< lltllNir ttirntfMtfftf CUrt* fcteO ftWIHWllf MM/ tfVyWQ »is€Ot Bare-eer ifpemaws Clade T r L • «« «p Rww Whlte-manUf mrt-anurus Clade Vxc tmianurut £ ****** Orarvpe-ieo m»rc*i Clade v<9 nmAM §mvui SADDLE-BACK TAMARINS Sapuimrs fuSCICOMi WHITE -MOUTH TAMARINS S»gtmit/s mgn colhs Clade a JHg n pft fcf n fit**** i § a i^/j wAH MUS TACHED TAMARINS S mysiax Clade m m pMtMu* 0 m etae Red "Cheated M- T. S. Clade Iduhri wtottrm - r momnd Emperor U , T S rmperarof Clade Figure 12. D istribu tions of all N eo tropical Tam arin Monkeys, genus SaguiflUS, com pared with those of all Amazonian Marmosets. 130 Marc G.M. van Roosmalen & Tomas van Roosmalen bleached taxa S. Uligeri and S. nigrifrons, and after crossing the upper Amazon River (where it is called Rio Maranon) northward into the progressively bleached taxa S. lagOUOtUS, S. fusCUS, and S. tri- partite, the latter three taxa being distributed north of the Amazon River in the Ecuadorian, Colombian, and Brazilian Amazon. Within the Black-mantle White-mouth Tam ar ins of the S. nigricollis Clade (Fig. 14) that is dis- tributed only north of the Amazon River in the Brazilian, Ecuadorian and Colombian Amazon, the nearest to archetypic saturated eumelanin taxon is S. nigricollis. It radiated northwestward and di- verged into the slightly bleached taxa S. graellsi and S. hernandezi. The S. nigricollis Clade is sympatric with the saddle-back tamarins of the taxa S. lagonotus, S. tripartitus and S. fuscus (Fig . 1 3 ) . However, they occupy different ecological niches and therefore can be seen traveling and foraging in mixed species associations, with the larger-sized black-mantle tamarins in the lead and staying higher up in the canopy of the terra firm e rain forest. Within the Emperor Mustached Tamarins of the S. imperator Clade both extant taxa are already progressively bleached, the grayish taxon S. Sllb- griscescens slightly more so than S. imperator. In the upper Rio Purus region there must exist a narrow contact zone between the two taxa along the southernmost open-end distribution of S. imperator. Within the Red-chested Mustached Tamarins of the S. labiatus Clade, Saguinus labiatus occupies the southernmost distribution and represents the nearest to archetypic taxon with a dark red chest and thin-lined white mustache. It radiated north of the Rio Ipixuna and diverged into the advanced orange- chested taxon S. rufiventer that has a m ore bleached white mustache and head-stripe. The third taxon of the S. labiatUS Clade is S. thomasi the precursor of which once must have traversed the Rio Solimoes. It m ight have been replaced later by S. illUStUS north of the Rio Solimoes as far west as the Rio Japura. Saguinus thomasi nowadays only occupies the lower Rios Solimoes /Japura interfluve. It represents the most progressively pheomelanin bleached taxon of the S. labiatUS Clade in its light orange-colored chest and the broad-lined triangular white mus- tache. Within the mustached tamarins of the S. mystax Clade, the more saturated eumelanin, nearest to archetypic form is represented by the taxon S. mystax that is distributed west of the Rio Jurua. After traversing the Rio Jurua, the Clade has radiated eastward while further bleaching along the pheomelanin pathway into the orange-crowned taxon S. pileatUS, and along the eumelanin pathway diverging directly from S. YYVyStaX into S. plutO. The latter taxon is overall more grayish and has a dis- tinctive albinotic spot around the base of the tail. In the lower Rios Jurua /Purus interflu ve we have sighted S. plutO ranging always in mixed-species association with the smaller saddle-back tam arin S. avilapiresi, with S. plutO always in the lead and S. avilapiresi rushing behind and below the group of S. plutO in the lower strata of high forest, always in a hurry feeding on S. plutO’s left-over food items. A hypothetical pathway of allopatric speciation, radiation and metachromic bleaching followed by the B are-face Tamarins of the S. midas, S. bicolor and S. geoffroyi sub-Clades m ay have had its origin in the Guianas (Fig. 15). An all-black, saturated eumelanin archetypic precursor of S. midas may once have traversed the lower Rio Amazonas and speciated allop atric ally into the black-handed taxon S. niger. Or vice-versa (archetypic black-handed S. niger may once have traversed the lower Rio Amazonas and speciated allopatrically into the red- handed S. midas). The same or another all-black precursor of S. midas may have traversed the Rio Negro and allopatrically speciated into the taxon S. inuStUS that is all-black with a white-mottled face. Saguinus inuStUS nowadays occupies the entire interfluve between the Rio Negro in the north, and the Rios Solimoes, Japura and Caqueta in the south. A founder-colony of a predecessor of S. inuStUS driven by the trend to allopatry may then have ventured from the taxon’s westernmost distribution into the NW Colombian Rio Magdalena basin. Once having inhabited the Rio Magdalena basin, it may have diverged along a pheomelanin pathway into the extant taxon S. leucopus that has a white- hairy facial circumference similar to S. inUStUS. Saguinus leucopus then may have radiated further into the progressively pheomelanin bleached, almost euchromic taxon S. OedipUS, and from there into the near-albinotic taxon S. geoffroyi that is distributed from extreme NW Colombia into Panama, as such the farthest away from the center of dispersion of the Bare-face Tam arin Clade. With respect to the three derived euchromic taxa of the S. bicolor Clade, as we have mentioned elsewhere, these taxa find themselves in the process of being On the origin of allopatric primate species 13 1 rigorously displaced from their respective territories by the now sympatric archetypic saturated eu- m elanin red-handed tam arin S. midas. All three taxa (i.e S. bicolor , S. martinsi, and S. ochraceus) find themselves pushed with the back against the untra- versable Rio Negro and/or Rio Amazonas (Fig. 15). At present, the red-handed tam arin S. midas is wrapping up the last stage of its range extension towards the south to the cost of all three Bare-face Tamarins of the S. bicolor subClade. This battle over a specific ecological (feeding) niche, in which two sympatric, closely related primate taxa are involved, will inevitably lead to the extinction of the most euchromic among the two, that is the Bare-face Tamarins of the S. bicolor sub-Clade: the taxa S. bicolor, S. martinsi, and S. ochraceus (Fig. 15). The eumelanin S. midas sub-Clade might have originated in the Guianas north of the watershed with the northeastern Amazon formed by the Tumac-Humac Mo un tains and the open wet savan- nas of Roraima and Para. A predecessor of the S. midas sub-Clade, perhaps the extant S. midas itself, once may have circumvented the watershed between the Guianas and Brazil by traversing the Parii Savanna, whereafter it may have penetrated far southwards into the northeastern quadrant of the Brazilian Amazon. We assume that before some vicariance took place this vast territory or a large part of it was inhabited by precursors of the closely related Bare-face Tamarins of the S. bicolor sub- Clade. Apparently, as the two sub-Clades do occupy the same ecological niche, (proto )-midas sub- sequently has displaced (proto)-bicolor over most of its former range. This battle is still being fought over between S. Iflidas and each taxon of the S. bicolor sub-Clade, but it seems to come close to its end. The process of replacement is accelerated by deforestation and other human disturbance such as road-building that has taken place north of the rap- idly expanding megacity of Manaus. This ongoing story clearly demonstrates interspecific intolerance in closely related territorial monkeys that occupy and exploit the same ecological niche. It inevitably leads to displacement, or sooner or later extermina- tion of the more progressively bleached (eu- chromic) taxon. This kind of replacements may take place after a geographic barrier has been success- fully overtaken by the more saturated eumelanin (more adaptive and/or aggressive?) of two related taxa. Or: after a vicariance has removed a hitherto gene-flow impeding geographic barrier inbetween the distributions of two or more closely related species. Vicariance (from Latin vicariuS) means a pro- cess by which the geographic range of an individual taxon, or an entire biota, is split into discontinuous parts by the formation of a physical barrier to gene flow or dispersion . Today, the S. bicolor sub-Clade only inhabits a 20-30 km narrow strip of terra firm e rain forest alongside the southernmost edge of the Pre-Cam- brian Guayanan Shield. The three bicolor taxa are so to speak pushed with the back against rivers that happen to be the widest and most difficult to traverse on the entire S ou th - A m eric an continent: the Rios Negro and Amazonas. The three extant taxa of Bare-face Tamarins each occupy what is called a “dead-end distribution”. The distribution of the half-brown, half- w hite taxon S. bicolor measures not m ore than 20-30 x 200 km, delineated in the west and south by the Rios Cuieiras, Negro, Am azonas, and Urubu. Bicolor’s neighbor to the east - the almost fully bleached, ochraceous colored taxon S. Ochraceus - occupies the interfluve between the Rios Urubu and Uatuma. To the east of its distribution, the pheo- m elanin, light orange-colored taxon S. martinsi occupies the lower interfluve between the Rios Uatuma and Nhamunda (Fig. 15). Disputedly, a now extinct precursor of the Bare-face S. bicolor sub-Clade that once ranged somewhere to the north of the Amazon River, may have driven the three extant taxa of the S. bicolor subClade - each at a different stage of metachromic bleaching - into the small interfluvial dead-end distributions, that they occupy today. The saturated eumelanin (blackish- brown) red-handed tamarin S. midas that later ex- panded its range to the south, is now simultaneously invading the three remaining adjacent interfluvial stronghold territories of the S. bicolor sub-Clade. A sharp-line contact zone drawn between S. midas and S. bicolor territory has been notified by us in the early 1990s to run at 28-30 km north of and parallel to the Negro and Amazon Rivers. While running a halfway house for orphaned mon- keys situated right at the edge of the contact zone, we have repeatedly witnessed different social groups of S. midas raiding resident family groups of S. bicolor. These incidents invariably ended up 132 Marc G.M. van Roosmalen & Tomas van Roosmalen 13 SADDLE-BACK TAMARINS Saguinus fusticollis Clade % f avilapiresi f. primrUvus QfJtiigeri f. wedddli (f) mclanoicucus f. ieucogonys fuscicottfs fm.) crandaili Af. lagonows cruzlimai t. nigrifrons ._$((■) trip annus 1 f, fuscus t mura BLACK-MANTLE WHITE-MOUTH TAMARINS Saguinus nigricoUis Clade n. nigricollis % n. gractlsi 0 n. hamandazi MUSTACHED TAMARINS S. mystax Clade m.mystax m. pileatus m. phito Red-chested M. T. S. labiatus Clade labiatus I. rufivanter I. thomasi Emperor M, T. S, imperator Clade imparmor i. subgrtsascens Figure 13. Distributions, allopatric speciation. radiation, and supposed pathways of metachromic bleaching in all known Saddle- back Tamarins of the Saguinus fuscicollis Clade. Figure 14. Idem, in the more robust, larger-sized Black-mantle White-mouth Tamarins of the S. nigvicollis Clade, the Emperor Mustached Tamarins of the S. iwiperatOV Clade, the Red- chested M ustached Tam arins of the S. IcibiutliS C lade, and the M ustached Tam arins of the S. tnystClX C lade. On the origin of allopatric primate species 133 BARE-FACE TAMARINS GLADE Saguinus gepffroyi Saguinus ocdipus Saguinus hucppuS • ' Saguinus bicolor Saguinus marlin si ocbraceus Saguinus maninsi maninsi 9 Saguinus midas # Saguinus nigpr Saguinus inustus 16 LION TAMARINS - Leontopithecus L. chrysopygus L. cbrysomelas L caissara i L rosaiia TRUE MARMOSETS OR OUISTIT1S - Catfithrix X C. jacchus |1C. panklllata .yt C. kuhfii C, gvoffroyi C. flaviccps C. aurita Figure 15. Distributions, allopatric speciation, radiation and supposed pathways of metachromic bleaching in all extant B are-face Tamarins that belong to the SagllinUS midciS, S. bicoloY, and S. geoffvoyi sub-Clades. Figure 16. Distributions, allopatric speciation. radiation and supposed pathways of metachromic bleaching in all known True (or Atlantic Forest) Marmosets (genus Cdllithrix ) and Lion Tamarins (genus LeOiltOpitheCUS) from SE Brazil. 134 Marc G.M. van Roosmalen & Tomas van Roosmalen in the defensive, less aggressive (more sensitive?) S. bicolor bitten to death. Now, about twenty years later, S. fflidciS has extended its range at least five km further to the south to the cost of S. bicolor oc- cupied territory. As S. midoS is more opportunistic and flexible in its habitat preferences - venturing also into secondary growth and edge habitats such as roadsides - it rapidly penetrates into S. bicolor territory, at some places (e.g., Ducke Reserve) already reaching the outskirts of Manaus. Running a rehabilitation center for orphaned monkeys, we sometimes received whole families of S. bicolor that were rescued from isolated pockets of forest in urbanized areas. After some time spent in quaren- taine, we used to put them in large cages built on poles in the middle of the rain forest about thirty km north of Manaus in an attempt to reintroduce the species where we assumed it had occurred not long before. One day before releasing a wild-caught social group of 8 S. bicolor, we found them all bitten to death inside the cage that was fenced with galvanized small-meshed wire. The only animal left alive in the cage was a wild adult *S. midos that apparently had not found back the little hole in the wire through which he and some other family mem- bers had entered the cage that very morning. On the other hand, a hand-tame S. lfliddS infant that we raised free around the compound at the time, one day was ‘kidnapped’ and adopted by the wild S. midas group that roam ed around in the project area. Within the True (Atlantic Forest) Marmosets or Ouistitis genus Ccillithrix we distinguish two monophyletic Clades: the Cd. penicillcitCl C lade and the Cd. OliritO Clade (Fig. 16). Within the first m onophyletic Clade w e consider Co. penicilloto the nearest to archetypic, most saturated eumelanin taxon that occupies the largest distribution (dark green area). From there, it radiated in northern direction and diverged into the overall progressively bleached taxon Co. jocchus that has fully albinotic ear-tufts. In eastern direction, from it derived and radiated away the taxa Co. kuhlU and Co. geojfroyi that are progressively bleached euchromic to al- binotic in their mantle and head parts (except the black ear-tufts). Their dead-end distributions are pressed against the A tlan tic coast. Interestingly, Co. kuhlifs range fully overlaps with that of LeOUtO- pithecus chrysomelos. The Co. aurita Clade has Co. aurita representing the nearest to archetypic, overall metachromic agouti taxon that ranges allopatric with the lion tamarins ( LeOUtopitheCUS ) in the Atlantic forest of SE Brazil. From there derived the near albinotic taxon Co. floviceps that occupies a small area in SE Minas Gerais, allopatric with Co. geojfroyi (M itterm eier et al., 2013). As for the Lion Tamarin genus LeontopitheCUS, we consider the overall saturated eumelanin, almost all-black taxon L. chrySOpygUS the nearest to ar- chetypic lion tamarin. From it derived in southeast- ern direction the taxon L. coissoro that followed a metachromic pathway of pheomelanin bleaching in its bright orange-colored dorsal parts while maintaining the saturated eumelanin black tail, arms, legs, mantle and head of L. chrySOpygUS. Its small range in coastal Parana State represents the southernmost distribution of any callitrichid. From L. chrySOpygUS derived in northeastern direction along a pathway of pheomelanin bleaching the two other taxa, L. chrysomelos and L. rosalio. Leonto- pithecus chrysomelos bleached in the orange colored lower arms and legs, and in the light orange to cream-white head and mantle maintaining the rest of its body saturated eumelanin. LeontopitheCUS WSolia, in turn, is evenly light orange-colored over its whole body, with the tail becoming almost al- binotic. Both taxa occupy small dead-end distribu- tions in the Atlantic forest along the coast of SE Brazil (Mittermeier et al., 2013). In a further attempt to falsify the principle of metachromic bleaching and the crucial role we believe it plays in allopatric speciation of (at least) Neotropical monkeys, we now will proceed to examine currently known distributions, allopatric speciation and radiation, and the pathways of metachromic bleaching supposedly followed in all other male-territorial Neotropical monkey genera (i.e., Callicebus, Saimiri, Cacojao , Chiropotes, Pithecia, Lagothrix, Ateles , Brachyteles, Alouatta, Cebus, Sapajus, and Aotus). Titi M onkeys of the genus Callicebus are strongly territorial in behavior, a family marking its territory vocally - a pair calling in duet, or a whole family calling in chorus. In the Amazon, a single taxon of the Collared Titi Col. torquOtUS Group may occur in sympatry with a single titi of any of the other Non-collared Titi cladistic Groups, once the former titis are only found high up in the canopy of primary terra firm e rain forest. Collared titis occupy a different, more frugivorous feeding niche than the titis that lack the white collar. The latter prefer the On the origin of allopatric primate species 135 lower strata and edges of terra firm e rain forest, secondary growth, and savanna forest, being overall more omnivorous in their diet that contains also young leaves and insects, in addition to pulpy fruits (H ershkovitz, 1988; Hershkovitz, 1 990; M itter- m eier et al., 2013). In figures 17-20, we show the distributions of all known Titi Monkeys genus CallicebuS. Within the titi monkeys five phylogenetic cladistic Groups or C lades are recognized: Ceil. peVSOnatUS (south- eastern Brazilian taxa), Cal. torquatUS { Amazonian collared taxa), Cal. moloch. Cal. ClipreUS and Cal. donacophilus (Amazonian non-collared taxa) (Van Roosmalen et al., 2002). Within each titi Clade the irreversible pathway of metachromic bleaching towards partly or fully albinotic, from saturated eumelanin and saturated pheomelanin fields to white or colorless, is clearly demonstrated. The farther radiated away from the prototypic agouti or saturated eumelanin (black or dark brown) taxon - Cal. melanochir in the Cal. personatus G ro u p , Cal. medemi in the Cal. torquatus Group, Cal. cineras- cens in the Cal. moloch Group, Cal. brunneus in the Cal. cupreus Group, and Cal. modestus in the Cal. donacophilus G roup - the more its pelage turns into orange, yellowish or cream to white, first in certain parts of the body, and eventually all over its coat. N ear-albinotic forms in dead-end distributions (e.g., Cal. pallescens, Rio Xingu titi, Rio Mam uni titi) are doomed to eventually go extinct, as meta- chromism with the trend to allopatry as the driving behavioral factor is an irreversible, initially seem- ingly non - adaptive evolutionary pattern in all territorial monkeys. As shown in the maps, in the Amazon all distributions of titis without a white collar are occupied by just a single taxon and are delineated by rivers that function as (for titis that cannot swim) strong geographic barriers. Narrow contact zones between adjacent interfluvial distribu- tions surely do exist, usually near the headwaters, but nowhere interbreeding or hybridization between the two neighboring taxa has been reported to take place. Our extensive primate surveys carried out throughout the entire Amazon Basin have revealed that, in general, a given monkey taxon looks phenoty pically identical throughout its entire range. In contact zones or across opposite banks of rivers that dem ographically separate two phy logenetically related taxa, we have noticed interspecific boundary conflicts and vocal battles to occur regularly, in particular performed by social groupings of titis, howling monkeys and spider monkeys. In at least one contact zone between two differently looking titis we have been able to perceive the ‘trend to allopatry’ put in motion in metachromic bleached individuals that were deviant from the commonly seen phenotype. At the far northeastern corner of the distribution of Hoffmann’s Titi Monkey Cal. hoffmannsi a small founder-population of an al- binotic all-cream w hite form, that we provisionally named the “Rio Mamuru titi”, apparently has been pushed into a dead-end distribution between the right bank of the lower Rio Mamuru, the for titis inhospitable varzeas (seasonally white-water inund- ated floodplain forest) along the right bank of the Rio Amazonas, and the parapatric distribution of Cal. hoffmannsi to the east and south as far as the lower Rio Tapajos (Fig. 17). Only mtDNA se- quences may determine what taxonomic status we should allocate to this new, fully euchromic taxon: ‘color morph’ or ‘taxon in the making’. A color m orph of Cal. hoffmannsi, a subspecies to be nam ed Cal. hoffmannsi mamuruensis, or a valid new species to be described as Cal. mamuruensisl As mentioned before, here our ecospecies concept could be applied in case the population has been confirmed to be allopatric and genetically isolated (not allowing any gene flow) from the taxon it derived from, or when the enclave population has successfully adapted to a different ecological niche - in this case turning itself into a varzea versus terra firm e rainforest habitat specialist. Our ecospecies concept (hereafter named ESC) in combination with the phylogenetic species concept (PSC) is, at least in the field, more practical, less arbitrary, and better defined, in particular when used for the purpose of species and biodiversity conservation. The ESC would put an end to the academic discussion about the arbitrary and controversial subspecies/race concept. As Groves (2001a; 2001b; 2004; 2005) points out: “ There is no official taxonomy ” .The numerous concepts as to what is and what is not a species are controversial, and every named species is itself nothing more than a hypothesis. Our understanding of the systematics of the primates is constantly growing, not only through the discovery of new species but also with new information brought to bear from diverse fields such as morphology, cyto- genetics, molecular genetics, paleontology, biogeo- 136 Marc G.M. van Roosmalen & Tomas van Roosmalen AMAZONIAN NON-COLLARED T1TI MONKEYS CALLICEBUS Callicebus moloch Clade C. cinerascens C. baptista C hoffmannsi C. moloch &C. vieiraiwC. bernhardi Callicebus cupreus Clade C, brunneus C. cupreus%C. caquetensis C caligaius € C- siephennashi C. aureipalatii dubius C. dlscolorGC. ornatus Callicebus donacophilus Clade C. modesws C. ollalaeM C. donacophilus C. sp nov. Chiquiianos oenanthe Chitjuiianen Titi Rit> Paraguay Caiiicut $p. /tov. Figure 17. Distributions, allopatric speciation, radiation and supposed pathways of metachromic bleaching in all known Amazonian N on-collared Titi Monkeys genus Callicebus. graphy, physiology and behaviour - contributing to test the hypothesis that a certain organism is a species distinct from another. Distinct in what sense? An individual is distinct, a population is distinct, but when and in what way is it a distinct species? Among the Titi Monkeys of the Cal. TTloloch cladistic Group (Fig. 17), the all-agouti dark-tailed taxon Cal. cinerascens, ranging along the east bank of the Rio Aripuana and between the right bank of the lower Rio Madeira and the left bank of the Rio Canuma, seems to represent the nearest to ar- chetypic, most original or ancestral titi from which all other tax a of the Cal. moloch Group have derived. Phy logeographically, the current central- southern Amazonian distribution of Cal. dneras- CenS is thought to represent the center of dispersion of the Cal. moloch Clade. In other words, the upper Aripuana region in Rondonia may be considered the cradle of Cal. moloch C lade’s evolution and disper- sion. From there, all taxa of the Cal. moloch Clade have diverged, radiating away in all (but southern) directions. A distinct metachromic trend to saturated pheomelanin (orange beard and sideburns) and al- binotic (cream to white beard, sideburns, tail and/or whole body) can be seen, which means that the most progressively bleached taxa that demograph- ically radiated the farthest away from the arche- type’s origin of dispersion tend to euchromic or albinotic (i.e.. Cal. moloch east of the Rio Tapajos, and Cal. hoffmannsi in the northernmost dead- end distribution delineated by the un traversable Amazon and Tapajos Rivers). The supposed meta- chromic pathway taken is as follows: Cal. dneras- CenS radiated first in northern direction, some founder-colony traversed the Parana do Uraria, followed the pheomelanin pathway, and diverged On the origin of allopatric primate species 137 COLLARED Till MONKEYS CALLICEBUS TORQUATUS CLADE Callicebus torquaws Subclade #C. r. torquaws C. t. purinus C. r. regulus Caflicebus luge ns Sub clade Figure 18. Distributions, allopatric speciation. radiation and supposed pathways of metachromic bleaching in all known Amazonian Collared Titi Monkeys of the Callicebus tOVquatUS Clade with two sub-C lades: Cal. tOVquatUS and Cal. lugetlS. into taxon Cal. baptista (which has dark orange- colored beard, sideburns, lower extremities, and belly). Radiating eastwards, it diverged into Cal. hoffinannsi (its forehead, beard, sideburns, hands, feet, and belly bleached light gray to cream -w hite). After Cal. hoffinannsi happens, A to traverse the Rio Tapajos, most likely where it is called Rio Juruena, it diverged into the advanced pheomelanin bleached to albinotic taxon Cal. moloch that now occupies a large distribution east of the Rio Tapajos and south of the Amazon River. Callicebus hoffinannsi also diverged along the upper course of the Rio Tapajos into the recently described taxon Cal. vieirai (ran- ging between the Rios Juruena and Teles Pires), which is near-albinotic. When a taxon is occupying a given interfluvial distribution delineated by hard to traverse river barriers, it has irreversibly changed its pelage or parts of its coat (e.g., beard, sideburns, ear-tufts, forehead, tail, hands, feet) following the eumelanin pathway from agouti or saturated eumelanin to al- binotic (cream or white), via black, brown, drab, and gray, and/or the pheomelanin pathway via red, orange, and yellow, or a combination of the two pathways in different parts of the body or coat. The trend to albinotic in the Cal. vnoloch Clade is completed near its northernmost dead-end distribu- tion in the all-cream to white new form that we happened to identify along the right bank of the Rio Mamuru (Fig. 21). It must have derived from dark- tailed but cream -bearded and -bellied Cal. hoff- mannsi. Following the trend to allopatry, this color morph (or ecospecies or ‘taxon in the making’?) is pushed with the back against the varzeas (white- water floodplain forests) and right bank of the untraversable Rio Amazonas. Callicebus moloch Clade’s westernmost distribution is represented by the advanced pheomelanin bleached (bright orange 138 Marc G.M. van Roosmalen & Tomas van Roosmalen belly, beard and sideburns) to albinotic (white fore- head, hands, feet and tail tip) taxon Cal. bemhardi which, in turn, is pushed with the back against the also u ntraversable Madeira River. Among the Titi Monkeys of the Cal. ClipreUS cladistic Group (Fig. 22), we consider Cal. brutl- neilS the nearest to archetypic taxon. Centrally dis- tributed, the overall agouti colored taxon Cal. brUYUlCUS radiated in northwestern direction via the progressively pheomelanin bleached taxa Cal. du- biliS and Cal. discolor into the most pheomelanin bleached (light orange tail base, beard, sideburns, belly and inner limbs) to albinotic (snowwhite tail, hands, feet and front/blaze) white-fronted taxon Cal. OmatUS that is distributed north of the Amazon in the Colombian Amazon. From Cal. brUYlYieiiS southwards diverged the advanced pheomelanin bleached taxon Cal. aureipalatii in the Clade’s Titi Monkeys genus CaUicebus Schematic Map of the Distribution of the eastern Brazilian taxa CaUicebus personatus Clade ATLANTIC OCEAN Figure 19. Schematic map of the d istribu tion s of the SE Brazilian or Atlantic Forest Titi Monkeys of the CaUicebus personatus Clade, which are separated by (for these taxa) untraversable rivers. On the origin of allopatric primate species 139 Figure 20. Distributions, allopatric speciation, radiation and supposed pathways of metachromic bleaching in all known SE Brazilian or Atlantic Forest Titi M onkeys of the Ctllliccbus pCTSOnCltUS Clade . southernmost distribution (the Bolivian Amazon between the Rios M adre de Dios and Beni). From Cal. brunneus radiated away in northern direction first the slightly pheomelanin bleached taxon Cal. CUpreuS that now occupies a large interfluvial area west of the Rio Purus and south of the Rio S olim oes.Afteran ancestral founder-colony of Cal. CUpreUS managed to traverse the Rio Purus to the east, it diverged into the advanced pheomelanin bleached all-orange, but white-tailed taxa Cal. Ca- ligatus and Cal. stephennashi in the north eas tern - most dead-end part of Cal. CUpreUS Group’s distribution, as the Rio Madeira represents the second strongest riverine barrier on the South- American continent. Last but not least, also from Cal. CUpreuS derived in northwestern direction the recently described, advanced pheomelanin bleached taxon Cal. caquetensis that at present occupies a small, not yet fully identified area north of the Amazon and Caqueta Rivers in the Colombian lowland Amazon, allopatric with and south of the distribution of the white-fronted titi Cal. OTUatUS (Fig. 17). Among the Titi Monkeys of the Cal. donaCO- philuS cladistic Group (Fig. 17), we consider the overall agouti-colored taxon Cal. modestUS the most original, nearest to archetypic taxon. It occu- pies the Clade’s northernmost distribution deline- ated by the Rios Beni and Mamore. It radiated southwards into the slightly pheomelanin bleached orange-brown taxon ollalae. Following a eumelanin bleaching pathway. Cal. modestUS also radiated in southeastern direction, first into the near-albinotic taxon Cal. donacophilus, and from there into the fully albinotic taxon Cal. pallescens. The latter nowadays occupies the southernmost dead-end 140 Marc G.M. van Roosmalen & Tomas van Roosmalen Figure 21. Distributions of CallicebllS baptista, Ca. Ho ffniCUVlS /, and the “Rio Mamuru titi” - the latter perhaps to be con- sidered a new taxon or one ‘in the making’. This satellite image shows the location of an enclave population of fully albinotic titi monkeys that we have found to exist along the right bank of the Rio M amuru. This p op ulatio n is on the verge of extinction as it is pushed with the back against for titis inhospitable habitat - the seasonally inundated floodplain forest (varzea) along the Rio Amazonas and the outskirts of the rapidly expanding town of Pari n tins in the north, and lands occupied by Ca. hoff- mannsi stretching to the east as far as the Rio Tapajos. The species Ca. baptista belonging to the Ca. moloch C lade originally ranged only north of Parana do Canuma, P. do U raria and P. do Ramos, east of the lower Rio Madeira, south of the Rio Amazonas and west of the Parana do Ramos. South of this narrow distribution evolved the species Ca. hofftnannsi, which occupies a large distribution between Rio Canuma in the west, Rio Tapajos in the east, and Rio Amazonas in the north, east of Parana do Ramos and Rio Mamuru. Baptist's Titi is much more color ful being dark to bright red on the ventral parts and lower limbs, having a red beard and red sideburns, whereas the rest of its body is grayish to blackish agouti. Hoffmann’s Titi is basically two-colored grayish and y ello w ish - w h ite to almost white, its sideburns and beard being light cream-white. H owever, we spotted the Ca. baptista titis also along the west bank of the Rio Uira-Curupa, hence it once must have tra- versed the Parana do Ramos west of the town of Parintin s, form ing an enclave population there after it displaced Hoffmann’s titis from the interfluve delineated by the lower Rio Uira-Curupa and Rio Andira. We also spotted advanced metachromic bleached, near-albinotic, pale yellowish to all-white ‘color morphs’ being phenotypically most related to Ca. hofftnannsi along the Rio Mamuru, the next river to the east, and classic yellowish- white and gray Ca. hoffmannsi with black tails along both banks of the middle and upper Rio Andira. These observations may confirm a case of what is called parapatry. The tw o valid species Ca. hofftnannsi and Ca. baptista that are allopatric for the greater part of their distributions - phy lo- geographically separated from one anotherby un traversable wa ter bodies - exclude one another where Ca. baptista happened to traverse a riverine barrier and subsequently replaced the local Ca.hofftnatinsi population. There, both taxa live parapatric, meaning in adjacent ‘patrias’ not separated by geographic barriers, where gene flow in theory is possible, but in reality does not occur. A plausible explanation would be that the two taxa have already diverged too far from one another. One could only speculate about the future of the fully albinotic form seen along the right bank of the Rio Mamuru. It may rep re sent a founder-colony or population of metachromic progressively bleached individuals that have been driven into parapatry by the Ca. hofftnannsi populations found to the east and south of R io M am uru as far as the R io Tapajos. The Rio Mamuru titis eventually might go extinct, unless they manage to adapt to (for titis) inappropriate habitat - the extensive varzeas along the right bank of the Rio Amazonas. If the founder-colony, following the trend to allopatry, would successfully adapt to the ecological niche of varzea, then a new taxon could derive from Ca. hofftnannsi. Through inbreeding, the currently adopted euchromic coat coloration would stabilize phenotypically across the entire population of that new taxon in a relatively short period of time. The hypothetical evolutionary path would then go from a metachromic fully bleached, near-albinotic color morph in a dead-end distribution to a new taxon belonging to the monophyletic Ca. moloch C lade . In that case, we would have to name the Rio Mamuru Titi Monkey Ca. tnattUlVUensis. On the origin of allopatric primate species 141 distribution of the Cal. donaCOphilus Clad e , penet- rating far into the arid Chaco of Paraguay and the pampas ofArgentina.A new species of titi, recently collected by the Brazilian ornithologist Marcelo Vasconcellos in the Chiquitanos area along the Rio Paraguay in the Pantanal of Mato Grosso do Sul (for which taxon we identified the holotype in the zoological collection of the AMNH, in 1977 collec- ted by George Schaller and m isidentified as Cal. donaCOphilus) , represents the easternmost distrib- uted taxon of the Cal. donaCOphilus C lade . Except for its dark gray ears (white in Cal. donaCOphilus), the Chiquitanos titi is overall more pheomelanin bleached towards albinotic than Cal. donaCOphilus , but less so compared to Cal. pallescens. Further- more, from Cal. modestUS derived in northwestern direction the advanced pheomelanin bleached near- albinotic taxon Cal. Oenanthe that is nowadays found isolated in a small area in the east-Peru vian Amazon, south of the Rio Maranon. Among the Collared Titis of the Cal. torquatUS cladistic Group (Fig. 18), we consider the saturated eumelanin black-handed taxon Cal. medemi w ith the westernmost distribution north of the Amazon River the nearest to archetypic form from which derived the all-black but yellow -handed titi from the southbank of the Rio Negro - a newly identified, as yet to be described taxon - and from that taxon derived the all-black, dorsally slightly reddish- tinged taxon Cal. lugenS with the northernmost distribution of the Cal. lugeYlS sub-Clade. From archetypic Cal. medemi south of the Rio Caqueta derived, first in eastern direction the dorsally pheo- melanin bleached taxon Cal. lucifer. Some ancestral founder-colony of the new Rio Negro southbank species then must have managed to traverse the lower Rio Solimoes somewhere between the mouth of the Rio Purus and that of the Rio Madeira. From there, collared titis could radiate away back in western direction, though south of the Amazon River, into the further pheomelanin bleached, overall reddish-brown colored white-handed taxa Cal. torquatus and Cal. purinus , and, after tra- versing the Rio Jurua, into the advanced pheo- melanin bleached red-handed red-fronted taxon Cal. regulus. Within the SE Brazilian Titi Monkeys of the Cal. personatUS C lade (Figs. 19-20) the nearest to archetypic, most saturated eumelanin taxon is Ca. melanochir. It ranges along the Atlantic coast south of the Rio Paraguagu in the center of dispersion of the personatus Clade. From Cal. melanochir de- rived in northern direction along the pheomelanin pathway the advanced pheomelanin bleached (all- orange colored) taxon Cal. barbarabrownae, and, in a small dead-end distribution delineated by the untraversable lower Rio Sao Francisco in the north and the Atlantic Ocean in the east derived the almost fully bleached, near-albinotic taxon Cal. COimbrai. Radiating in southern direction, ancestral Cal. melanochir diverged along the eumelanin pathway into the orange-tailed, but overall dark brown-colored taxon Cal. nigvifrons , and along the pheomelanin pathway into the advanced pheo- melanin bleached, all-orange colored and near-al- binotic taxon Cal. personatus. Within the Squirrel Monkeys genus Saimiri (Fig. 22), we phylogeographically distinguish two monophyletic Clades: Sa. SCiureus - including the C entral-A m erican Sa. oerstedii sub-Clade - and Sa. boliviensis - including the Bare - ear Sa. UStUS sub- Clade (H ershkovitz, 1 984). It is inferred that the genus Saimiri evolved relatively recently, with crown lineages diverging as late as the Pleistocene (ca. 1.5 MYA) and other major Clades diverging between 0. 9-1.1 MYA. Concurring with Chiou et al. (2011), we include Sa. Oerstedii in the mono- phyletic Sa. SciureUS Clade that originated in the Guianas. North of the Amazon, it radiated in west- ern direction and diverged first into Sa. Cassiquiar- ensis, a taxon that is nowadays distributed across the entire Rio Negro basin, its distribution in the south delineated by the Rio Japura/C aqueta and the lower Rio Solimoes. From Sa. Cassiquiarensis di- verged in northern direction the advanced bleached, least colorful taxon Sa. albigena that ranges allo- patric (north of the Rio Guaviare) in the southwest- ernmost part of the Rio Orinoco basin. In concurrence with Chiou et al. (2011), who found evidence for monophyly in the Sa. SciureUS and Sa. oerstedii Groups, we suggest that from Sa. albigena or some ancestral precursor of it derived and radi- ated away in northwestern direction the advanced pheomelanin bleached taxa Sa. oerstedii and Sa. citrinellus. These now range in Sa. SciureUS C lade’s disjunct northw esternm ost dead-end distribution - along the Pacific coast of Panama and Costa Rica. Along a different metachromic pathway derived from Sa. Cassiquiarensis in southern direction the advanced bleached taxon Sa. macrodon. Its distri- 142 Marc G.M. van Roosmalen & Tomas van Roosmalen bution is delineated by the Rios Guaviare and Apa- poris in Colombia, and the Rio Japura in Brazil, and south of the Amazon by the upper Rio Maranon in the west, and the Rio Jurua in the east. Within its large distribution, Sd. IflClCwdon is excluded from the Rios H uallaga/U cay ali interfluve in the Per- uvian Amazon that is occupied by Sd. pevuviensis. In the Guianas, Sd. SCiureiiS once managed to tra- verse the lower Amazon River to the south. As it is a riverbank marsh and mangrove forest specialist, Sd. SCiureiiS must have colonized the south bank of the Amazon after reaching it on floating islands covered with varzea or mangrove vegetation. From Sd. SCiureuS south of the Amazon subsequently de- rived the recently described, advanced bleached near-albinotic taxon Sd. COllillsi that is confined to M arajo Island - the Sd. SCiureuS Clade’s eastern- most dead-end distribution delineated by the At- lantic Ocean, and the Amazon and Para Rivers. The second monophyletic Clade of Squirrel Monkeys, the Sd. boliviensis Clade, has origin- ated in the extensive white-water floodplain forest (varzea) near the confluence of the Japura and Solimoes Rivers. The lower Japura/Solim oes inter- fluve does not contain any terra firme. It is season- ally flooded over 6-8 months. Here lives the nearest to archetypic, saturated eumelanin taxon of the Sd. boliviensis Clade, Sa.vanzolinii. It is overall agouti and black colored, representing the only extant squirrel monkey with an all-black tail. A somewhat bleached Sd. vanzolinH founder-colony once must have reached (swimming or on a floating varzea is- land) the south bank of the Rio Solimoes east of its confluence with the Rio Jurua. There evolved from it the somewhat pheomelanin bleached, orange to yellowish taxon Sd. boliviensis. It then occupied east of the Rio Jurua the entire area delineated by the lower Purus, upper Madeira and Guapore S . S. coliins SAIMIRI Saimiri sciureus Clade S sciureus S, cassiquiarensis #S, afbigena&S. macrodon Saimiri oersted// Clade oersredii eitrinellus Saimiri boliviensis Clade ^ S. vanzoiinii #S. boliviensis peruviensis S. usws Figure 22. Distributions, allopatric speciation, radiation and supposed pathways of nietachromic bleaching in all known Squirrel Monkeys genus Saimiri divided up in the S. SCiureUS and S. boliviensis C lad e . On the origin of allopatric primate species 143 Rivers, whereas in the south it extended its range far into the Peruvian and Bolivian Amazon. After a founder-colony of somewhat bleached Set. bolivien- sis happened to traverse the easternmost river bar- rier, it diverged into Sa. UStUS - the least colorful, most eumelanin bleached taxon of the Sa. bolivien- sis C lade. Its distribution is confined by the Amazon River in the north, the Rio Xingu in the east (sep- arating the distributions of Sa. UStUS and Sa. SciureilS) , and the Rio Guapore in the south. Fur- thermore, from boliviensis in its southw esternm ost distribution in the Peruvian Amazon derived the progressively pheomelanin bleached near-albinotic, most colorful taxon Sa. peruviensis. It occupies a dead-end distribution - the interfluve delineated by the Rios Huallaga and Ucayal i - as it is surrounded by Sa. macrodon occupied territory. Historically followed metachromic and phylo- geographic pathways, intraspecifically pushed ahead by the trend to allopatry and the principle of metachromic bleaching, within a genus or mono- phyletic Group of primates may be traced back most expressively, when we examine the distribu- tions, speciation and radiation of all extant Uakari Monkeys genus CacajaO (Figs. 23, 24). This ex- clusively Amazonian genus contains two mono- phyletic Groups or Clades: the Black-headed Uakaris of the Cac. melanocephalus G roup , and the Bald-headed Uakaris of the Cac. Calvus Group (H ershkovitz, 1 987a). Among Uakaris, the sup- posedly nearest to archetypic (prototypic) ancestral form is represented by the extant Black-headed Uakaris, more in particular by the saturated eu- melanin, all-black taxon Cac. ayresi - the north- easternmost distributed among all Uakaris. Uakaris are the only monkeys in the Neotropics that lost a functional tail. All other genera evolved either a long pendulous, short-hairy to bushy tail that is in the first place designed to use for balance while moving through the tree tops; or, a long pre- hensile tail that is used as a fifth limb during vertical climbing and walking on top of or brachiating un- derneath twigs and thin branches in the periphery of tree tops (where the fruits are hanging). Only after observing Black-headed Uakaris Cac. hosomi in the wild along the Rio Cauaburi and in Pico da Neblina National Park, we came to understand why the region drained by the Rio Negro has to be con- sidered the center of dispersion for all uakaris, in other words the cradle of evolution of the genus CacajaO. Simultaneously, we came to understand the very reason why uakaris have lost a functional tail, whereas in all other canopy-dwelling monkeys from the Amazon it seems to be a fifth limb of vital im portance. Across the entire upper Rio Negro basin the type of vegetation that dominates the landscape is a very impoverished sort of thin-stemmed savanna forest. It stands on poorly drained, highly acidic white- sand soils that are deposited on top of an imper- meable, several meters thick layer of coarse rounded pebbles. This type of forest is called “caatinga-do-Rio-Negro”, for it resembles much the arid dry seasonally deciduous vegetation in large parts of the Brazilian northeast. It seasonally floods during the long rainy season, but also throughout the year on a daily base during heavy rainstorms. Phy siognom ically, this forest type resembles two- storey mangrove forest, as most of its trees use pneum atophores (aerial roots) and stilt-roots to cope with frequent flooding conditions. Phytoso- ciologically, the ‘caatinga-do-Rio-Negro’ is dom- inated by trees belonging to families like Euphorbiaceae and Apocynaceae, known for their often toxic latex and plant parts, most in particular full-grown seeds. Surprisingly, this forest lacks hem i-epiphy tic climbing shrubs, vines, and twiners. Over geological times Black-headed uakaris seem to have co-evolved with this natural environment through specializing themselves in the depredation of immature seeds. From early maturation on, the seeds are often loaded with toxic alkaloids and secondary compounds. Uakaris have guts that are specially adapted to neutralize these toxins. Their canines are oversized and wedge-shaped with razorblade sharp edges, as such adapted to open up the toughest-husked fruits and kernels (endocarps) around. Their incisors are procumbent and used to scoop out the seed content (endosperm) from any endocarp or pericarp. Uakaris are full-fashioned seed predators to such length that, if one offers a uakari a juicy pear or apple, the monkey will in- stantly bite the pulpy pome in half with its powerful canines. Then, it will pick the tiny seeds from the central part, discard the pulp, and delicately split the tiny seeds one by one with their canines. In the end, it has its procumbent incisors scoop out the en- dosperm from the seed coat. Black-headed uakaris do occupy very large home ranges. They restlessly travel or forage in very large multi-male dominated 144 Marc G.M. van Roosmalen & Tomas van Roosmalen social groups that may contain over two-hundred monkeys. Since their preferred habitat ‘caatinga- do-Rio-N egro’ basically lacks climbing shrubs, the tree tops are not interlinked by vines, twiners and climbing hem i-epiphytes as they are in primary terra firme rain forest elsewhere in the Amazon. By lack of a walkway through the tree tops, Black- headed uakaris co-evolutionarily have adapted to this ancient impoverished, physiognom ically discontinuous and frequently inundated forest type by developing the locomotor pattern of so-called ‘vertical clinging and leaping’. A traveling or for- aging troop of Black-headed uakaris much re- sembles Madagascar indris, Indri indri (Gmelin, 1788) that also make enormous leaps, catapulting themselves for - and upwards by means of their strong muscular upper legs. Like indris in Mada- gascar, black-headed uakaris lost most of a func- tional tail while adapting to this type of locomotion. The few cm long tail provided with a tuft is only used for intragroup communication. Black-headed uakaris can curl it upwards and wave it sideways like dogs would do with a largely amputated tail. Black-headed uakaris of the species Cac. ho- SOtfli and CdC. Ciyresi , distributed north of the Rio Negro, east of the Cassiquiare and west of the Rio Demeni, and Cac. melanOCephaluS from south of the Rio Orinoco, west of the Cassiquiare and north of the lower Rio Solimoes and Rio Japura /Rio Caqueta, have a pitch-black face, a black, forward directed hair-tuft on the forehead, and a short black- ish, red or orange-tinged tail. Black-headed uakaris from the Rio Iqana basin, being distributed in - between the upper Rio Orinoco and the lower Rio Uaupes, show a black upper back and pheomelanin bleached, orange to blond bleached lower back. Perhaps, for that reason they should be taxonom- ically treated as a valid species (we here suggest Cac. ouakary). The Bald-headed Uakaris of the Cac. calvus cladistic Group, which range south of the Amazon/ Solimoes and Japura Rivers, have a bald head, bright-red bare face, blue-gray eyes, a shaggy pheo- melanin bleached, near-alb inotic coat, and a rudi- mentary tail that is shorter and even less functional than the tailofBlack-headed U akaris (Figs. 23, 24). The Cac. CalvUS Group contains five taxa which according to our phylogenetic ecospecies concept (ESC) should be all given valid species status: 1) CaC. CalvUS living exclusively in the white-water floodplain forests (varzeas) between the lower Japura and Solimoes Rivers, being cream-white with pheomelanin bleached, orange-brown ventral parts; 2) Cac. UOVaesi occuring in disjunct pockets along both banks of the lower and middle Rio Jurua as far upstream as its confluence with the Rio Ta- rauaca, its coat being pheomelanin bleached, orange brown-colored, but albinotic from the back of the head to mid-dorsum; 3) Cac. rubicundus, the pheo- melanin bleached, bright orange-colored (except for the albinotic cream-white back of the head and neck) bald-headed Cac. uakaris that occurs in the white-water floodplain forests (varzeas) along the left bank of the upper Rio Solimoes in the central- westernmost Brazilian Amazon; 4) Cac. UCayalU, its coat overall saturated pheomelanin, dark brown to orangish colored, ranging in the Peruvian Amazon along the right bank of the Ucayali River in the white-water inundated floodplain forest (varzea) as well as adjacent terra firme rain forest; 5) a form newly identified by us in the year 2000, its coat near-albinotic, advanced euchromic bleached to all-white. We provisionally name this new taxon the “Rio Pauini Bald-headed Uakari” CacajaO sp., for it is only found in the varzeas along the upper Rio Pauini, a left-bank tributary of the Rio Purus (Figs. 23, 24). The 1 trend to allopatry in metachromic varieties of sociable, but territorial primates' applies to the evolutionary path along which a certain primate race, species, monophyletic clade, or genus has ex- tended its geographic range in the geological past. As a founder - colony or - population at the outer limit of a taxon’s current range represents an ex- tremely narrow gene pool, through inbreeding certain phenotypic characters like partial depilation of the skin, or skin/coat coloration will be rein- forced in the beginning and therefore advance more rapidly. Through the process of metachromism (= evol- utionary change in tegumentary or hair/skin color- ation), with the ‘trend to allopatry’ in metachromic bleached individuals as the principal behavioral driving force, speciation, radiation, and phylogeo- graphy can be plausibly retraced and explained for in all extant N eotropical prim ates. A ccording to the principle of metachromic bleaching, primate taxa at the base of a phylogenetic tree or clade being the nearest to archetypic, prototypic, primitive, or ori- ginal, in general are agouti or saturated eumelanin, On the origin of allopatric primate species 145 UAKARIS CACAJAO Schematic Map of the Distribution of melanocephalus and calvus C lades Orinoco Cassiqulare Dcmoni Pheomelanln pathway Caquotd Uaupcs Branco Amazonas Amazonas Soiimdes Madeira Jurutf Maranon Pauini Javan' Pheometanin to albino Ucayali UAKARIS Cacajao melanocephaius Glade C. ay rest C.hosomi "|-C, melanocephaius Cacajao calvus Glade 0C. uc ay alii ^C. rubicundus C, calvus C. novaesi kC. sp. nov. Rio Pauinf i £2^3^ 9 \ \ V ^ ury> -(jr i w 1 23 24 Figure 23. Schematic map of the distributions of U akari Monkeys of the CaCCljaO inelcilWCephciluS and C. CCllvUS C lades div ided up by (for them ) un traversable rivers . F igure 24 . D istribu tion s, allopatric speciation. rad iation and supposed pa th ways of metachromic bleaching in all known Uakari Monkeys genus Co.CO.jciO. 146 Marc G.M. van Roosmalen & Tomas van Roosmalen which means the least colorful, agouti, black, or dark brown colored. Among Uakari Monkeys genus Cacajao, the origin or center of dispersion is sup- posed to be located in the northeasternm ost part of the Brazilian Amazon, south of the watershed between the Rio Negro and Rio Orinoco basins, an area delineated by the Rios Demeni and Araca (Figs. 23, 24; Fig. 27). Within this interfluve the landscape is dominated by ‘c aating a-do -R io - Negro’, the most impoverished habitat type ima- ginable, but preferred by uakaris of the CciC. melci- nocephalus Clade. Here lives the saturated eu- melanin, least bleached taxon of the Black-headed Uakaris, the recently described CciC. ayresi (B oubli et al., 2008). Its coat is all-black and dark-brown colored. It may well represent the proto- or ar- chetypic uakari from which all other uakaris have derived. From CciC. ay resi in western direction first diverged along the pheomelanin pathway taxon CciC. hosomi. It is distributed between the Rio Marauia, the upper Rio Negro, and the Cassiquiare Channel (we have confirmed its presence in Pico da Neblina National Park and along both banks of the Rio Cauaburi). After an ancestral founder-colony traversed the Rio Cassiquiare - the channel that connects the Rio Negro basin with that of the Rio Orinoco in Venezuela - CciC. IlOSOlfli diverged into an intermediately pheomelanin bleached taxon that differs from classic CciC. vnelciYlOCephaluS in the black shoulders, dark-red legs and tail. If this phenotype, which is thought to represent a color morph of Cac. melanocephalus, turns out to occur throughout the entire distribution delineated in the north by the Rio Orinoco and in the south by the Rio Uaupes, one should consider it a new taxon to be named the “Rio Igana Black-headed Uakari” Cac. Oliakary. After an ancestral founder- colony of the latter managed to traverse the Rio Uaupes, it has diverged into the progressively pheomelanin bleached blond- backed black-headed uakari taxon Cac. melanocephalus. Subsequently, blond-backed Cac. melanocephalus have occupied the entire interfluve south of the Rio Negro, eastwards as far as Archipelago de Anavilhanas located about forty km west of Manaus, and to the west far into the Colombian Amazon, and south as far as the north bank of the Rio Japura (Rio Caqueta in Colombia). We suppose that once upon a time a founder-colony of slightly bald-headed, advanced pheomelanin bleached ancestral Cac. melanocephalus , being pushed out of its westernmost dead-end distribution in the Colombian Amazon, may have managed to traverse the upper reaches of the Rio Caqueta. It then could extend its range southwards, eventually reaching the Rio M arahon (as the upper Amazon River is called in Peru). A fo under - colony of an advanced pheomelanin bleached, bald-headed ancestral form must then have traversed the Rio Ucayali. It subsequently occupied terra firm e and varzea forests in the interfluve between the Rio Ucayali in the west, the Rio Maranon in the north, and the Rio Javan in the east. Nowadays, this inter- fluve is inhabited by the bald-headed dark reddish- brown taxon Cac. UC ay alii that belongs to the bald-headed Cac. CalvUS C lade. D isjunct from Cac. UCayalii's distribution and ranging farther to the east derived taxon Cac. rubicundus , a progressively pheomelanin bleached bright- orange colored bald- headed uakari. It is fully adapted to varzea habitat found in abundance along the left bank of the upper Rio Solim oes. From Cac. rubicundus going farther eastwards, but disjunct from its distribution, along the same (left) bank of the S olim oes/A m azon River the almost fully albinotic taxon Cac. CalvUS is found. It fully adapted to white - water inundated floodplain forest (varzea) - the only available habitat in this for the Cac. CalvUS Clade dead-end distribution situated inbetween the banks of the Japura and Solimoes Rivers. Directly from Cac. rubicundus to the south of Cac. calvus ' distribution derived the bald-headed taxon Cac. UOVaesi that ranges along both banks of the Rio Jurua as far south as the confluence with the Rio Tarauaca and Rio Envira. This taxon is near-albinotic from the back of the head to beyond the mid-dorsum, and progressively pheomelanin bleached light orange- brown on the lateral and ventral parts of the body. It ranges in the varzeas of the floodplain, but we have also spotted large troops foraging for immature seeds in the adjacent terra firm e rain forest. In 2000, we identified a fifth taxon of bald- headed uakari, the completely white, fully al- binotic taxon that we named “Rio Pauini Bald- headed U akari” Cacajao sp. It lives along the south bank of the Rio Pauini, a left-bank tributary of the upper Rio Purus. It represents the southern- most distributed and the farthest pheomelanin bleached most albinotic taxon of all extant uakaris. It lacks the pheomelanin orange-brown to orange On the origin of allopatric primate species 147 ventral parts seen in the other near-albinotic taxa Cac. novaesi and Cac. calvus (Figs. 23, 24). Analyzing metachromic skin and coat characters as linear and irreversible progressions within Neo- tropical primate genera and their monophyletic Clades does add substantially to the reconstruction of bio geographic divergence events and phylo- genetic relationships over a wide range of Neo- tropical primate taxa, in particular those that defend their living space or ecological (feeding) niche through male-dominated, hierarchically organized societies. So it does to the Bearded Sakis genus Chiropotes (Figs. 25, 26) even if we have confirm ed in the field that social groups of (at least) the Guianan taxon Ch. SagulatUS do freely fuse and fission on a regular base with neighboring social groups. The genus Chiropotes clearly shows sexual dimorphism in the larger, more robust males that also grow bigger beards and frontal hair lobes on their heads (H ershkovitz, 1 985). During foraging and resting, a large social group of bearded sakis, similar to woolly monkeys, consists of several polygamous dominant males each taking care of his ‘harem’. The high-ranking males tend to stick to the center of the foraging troop, whereas lower ranking males with or without harems are pushed closer to the periphery of the foraging troop. This way, adult males do avoid confrontations, for their impress- ively large wedge-shaped canines designed to crack hard-husked fruits and kernels in order to get to the seed pulp would be lethal if used in fights. But ad- olescent, subadult, and, we assume, also behavior- ally or pheno typically deviant individual males may well be pushed into the periphery of the foraging and ranging troop. More than once, we have encountered a solitary male, or a couple of males traveling at high speed through the canopy in an apparently fixed direction, leaving us no means to determine if these monkeys only temporarily had lost contact with the troop, or if they were expelled from the parental troop, or if they were representing subtly deviant young males that had been forced to leave the pack and search for new living grounds somewhere beyond the limits of the group’s home range. Only through long-term field studies one would be able to obtain clear answers to this sort of questions. Within the Bearded Sakis genus Chiropotes we distinguish two monophyletic Groups: the Ch. SatanaS and the monotypic Ch. albinaSUS Group. The Ch. SatanaS Clade consists of five taxa, among which the nominate species Ch. SatanaS represents the saturated eumelanin, all-black, nearest to ar- chetypic bearded saki. Its distribution in the NE Brazilian state of Maranhao is assumed to represent the cradle of evolution or center of dispersion for the genus. An equally all-black form that we re- cently identified west of the headwaters of the Rio Xingu (e.g., Rios Ronuro, Batovi and Vonden Steinen) may either represent an enclave population that became disjunct from that of Ch. SatanaS (ranging east of the Rio Para/lower Rio Tocantins), or a new taxon of the Ch. SCltCMClS Clade that still has to be collected and described. From Ch. SatanaS diverged in western direction the slightly eumelanin bleached, overall light-brown colored taxon Ch. Utahicki. It occupies the entire interfluve delin- eated by the Rios A m azonas/A napu/Tocantins- Araguaia/X ingu . An ancestral founder-colony of somewhat pheomelanin bleached, red to orange- brown backed Ch. Utahicki once must have man- aged to traverse the lower Rio Amazonas, from which then derived taxon Ch. SaglilatUS. This species occupies the entire area north of the Amazon River and east of the Rio Branco, including most of the Guianas east of the Essequibo River. This taxon is absent from most ofAmapa state, French Guiana and also from a wide coastal belt of the Guianas. A founder-colony of ancestral sagulatus once must have traversed the Rio Branco and radiated in western direction diverging into the advanced eumelanin bleached taxon Ch. israelitCL. This species is characterized by the albinotic (white instead of pink) genitals and the light-grayish to brownish coat color of the trunk. Chiropotes israel- ita ranges west of the Rio Branco as far north as the Rro Orinoco in Venezuela. It seems to be parapatric with Black-headed Uakaris, as Chiropotes is a seed- predating terra firm e rainforest specialist, and Cac. ayvesi and Cac. hosomi are ‘caatinga-do-Rio- N egro ’ -habitat specialists. The Rios Marauia and Cauaburi seem to divide their distributions. O ur ex- tensive surveys in the Rios Demeni/Araca inter- fluve did not reveal the occurrence of Ch. israelita, as the landscape is dominated by ‘caatinga-do-Rio- Negro’ habitat (Figs. 26, 27). The second monophyletic Group of Bearded Sakis is that of monotypic Ch. albinaSUS. The Red- nosed Bearded Saki is very different from the Ch. SatanaS Clade, not just in metachromic sense. Its 148 Marc G.M. van Roosmalen & Tomas van Roosmalen utahicki 10 inches catvus ca!vus catvus ucayalii catvus novaesi catvus rubundus color form from Icana hosomi ayresi metanocephatus Figure 25. The hitherto recognized taxa ofBearded Sakis genus Chiwpotes (above) and Uakaris genus CciCCljciO (below ), all depicted in one plate (Courtesy of Stephen Nash). On the origin of allopatric primate species 149 BEARDED SAKIS CHIROPOTES Chiropotes satanas Clade Ch. satanas ^Ch. satanas Rio Xingu Ch. utahicki >Ch. sagulaws %Ch, israelita (was chiropotes) Chiropotes aibinasus Ch, aibinasus Figure 26. Distributions, allopatric speciation, radiation, and supposedly followed eumelanin pathways of metachromic bleaching in all known Bearded Sakis genus Chiropotes. vocalizations are very different, the beard and tail are shorter-haired, and the genitals of each gender are brightly red-colored as is the muzzle (the sci- entific name Ch. albinaSUS - Latin for “white nose”- relates to the taxonomist, who may never have seen the monkey he described alive. Furthermore, group size in Ch. albinaSUS is much larger than that of any of the taxa belonging to the Ch. satanas Clade, ranging on average from 30-80 individuals. W here Ch. albinuSUS occurs in sympatry with woolly mon- keys (i.e., west of the Rio Tapajos-Juruena, east of the Rio Madeira, and north of the Rio Ji-Parana), they are often seen in mixed species associations. Red-nosed saki groupings mixed with woolly mon- keys (Lagothrix cana), tufted capuchins ( Sapajus apella) and/or white-fronted slender capuchins ( Cebus unicolor ) m ay contain as m any as 150 mon- keys. In figure 28, we have depicted the distributions, allopatric speciation, radiation, and supposedly followed pathways of metachromic bleaching in all known Saki Monkeys genus Pitheda. Sakis occur exclusively in the rain forests of lowland Amazonia and the Guayanan Shield (Hershkovitz, 1987b; Mittermeier et al., 2013). Within the genus Pitheda we distinguish three monophyletic cladistic Groups: P. monachus, P. pitheda , and ^ hirsuta (Fig. 29). W ithin the P. monachus Clade allopatric specia- tion is thought to have followed evolutionary path- ways of metachromic bleaching with P. Monachus representing the nearest to archetypic precursor of all extant sakis. Both sexes have an overall satur- ated eumelanin, slightly bleached silky coat, except for the cream-white hands and feet. Taxon P. VUOn- achus ranges along both sides of the Amazon up- stream from its confluence with the Rios Jurua and 150 Marc G.M. van Roosmalen & Tomas van Roosmalen Figure 27. Map showing distributions of the Bearded Saki taxa Chiwpotes SClgulcitUS and C. israelita, and the parapatric Black-headed Uakaris that occur north of the Amazon and Negro Rivers. Japura, large rivers delineating its distribution in the east and north. The species is sexually dimorphic, not in size but in metachromic pelage characters of the head. Both sexes have a slightly bleached mask that is light brown in males and cream-white in fe- males. It surrounds a black face with yellow to cream eyebrows and malar stripes. Forehead and cheeks are covered with short, forward directed hairs resembling much that of members of the P. pithetia Group. From P. monachliS diverged in northwestern direction taxon P. milleri, supposedly after a metachromic deviant founder-colony of ancestral monachus traversed the Rio Caqueta. Pithecia milleri nowadays occupies a small part of the Colombian Amazon that is confined by the Rios Caguan and Putumayo. Both sexes are overall eu- melanin bleached, more so in females. The forehead is covered with long, forward directed hairs forming a kind of hood that is yellowish in males and cream- white in females. The black muzzle is contrasted with the advanced euchromic malar and lip stripes. From P. milleri derived the taxon P. napensis after a founder-colony of P. milleri traversed the Rio Putumayo in southern direction. Pithecia napen- sis occupies a small area in the Colombian and Ecuadorian Amazon delineated by the Rio Putu- mayo in the north and the Rio Napo in the south. In P. napensis both sexes are progressively pheo- melanin bleached in the yellowish to orange breast, more so in males that also differ in the silvery grayish lower part of a well-defined mask and in the albinotic hood. After a founder-colony of ancestral P. napensis once traversed the Rio Napo to the south, the progressively pheomelanin bleached taxon P. aequatorialis diverged. It occupies a large area in the Ecuadorian and Peruvian Amazon delin- On the origin of allopatric primate species 15 1 eated in the north by the Rio Napo and in the south by the Rio Tigre. Pitheda aequatorialis, in particu- lar in the metachromic characters of the male’s head (fully albinotic mask) and (orange) breast pelage, represents the most advanced pheomelanin bleached taxon in the P. TYlOYiachllS Clade. Its dead-end distri- bution at the end of the phylogeographic radiation of the P. monachliS Clade is confined at all but western (Andean Mountain range) sides by P. mOYl- CtchllS occupied territory. We may ponder about what would be the result of any hypothetical hy- bridization between P aequatorialis females and P. monachliS males at the contact zone that should exist in the species’ westernmost distribution. Even if the offspring would remain fertile, it would never result in parapatric speciation. In concurrence with our theory, deviant young males with metachromic genes from P. aequatorialis w ould be expelled by the dominant male(s) of the P. monachus parental group, back to P aequatorialis territory . Within the P. pitheda Clade we consider P. lotichiusi with the overall darkest agouti (in fe- males) and saturated eumelanin black (in males) pelage the nearest to archetypic taxon. This taxon is only found in the easternmost part of the inter- fluvial peninsula between the lower Solitudes and Negro Rivers, from opposite the city of Manaus as far west as the towns of Manacapuru and Novo Airao. In the past, P. lotichiusi m ay have occupied a much larger distribution, for no untraversable geographic barriers exist when going further west into the Rios S olim oes/N egro interfluve. If so, the P. pitheda Clade may have m onophyletically derived from the P. monachus Clade, when that ra- diated to the east. A founder-colony of slightly pheomelanin bleached ancestral P. monachus may SAKI MONKEYS PITH EC I A Pitheda pitheda Clade fl P paheciaQP, chrysocephala P. lotichiusi Pitheda monachus Clade P. monachus P. mitleri P. napcnsis Paoquaioriaiis Pitheda hirsuta Clade P. irrorata ^P. hirsuta P. vaniolinii P. albicans Figure 28. Distributions, allopatric speciation, radiation, and supposedly followed metachromic pathways of bleaching in all known Saki Monkeys genus Pithedci. 152 Marc G.M. van Roosmalen & Tomas van Roosmalen have traversed the lower Rio Japura and thereafter diverged into the allopatric taxon P. lothichiusi. The latter then extended its range to the east. During one of the late-Pleistocene glacials, when ocean levels dropped over up to 120 m, a founder-colony of P. loti Chius i could well have traversed the lower Rio Negro and then reached the north bank of the Amazon. This way, it may have diverged into the allopatric Golden-faced Saki taxon P. chryso- cephala. Nowadays,Golden-faced sakis range from the Rio Branco as far east as the Rio Trombetas. After a founder-colony of ancestral P. chrySO- cephala once traversed the Rio Trombetas, taxon P. pithecia may have diverged. Pithecia pithecia then expanded its range in northwestern direction across the states of Roraima, Para and Amapa, and across the Guianas into Venezuela as far west as the lower Rio Orinoco. It may have circumvented either side of the watershed formed by the Tumac Humac Mountains. Within the sexual dimorphic P. pithecia Clade, females are progressively pheo- melanin bleached orange to yellowish brown, whe- reas males are all-black with a progressively pheo- melanin bleached to albinotic mask. In the Brazilian taxa P. lotichiusi and P. chrysocephala the mask that consists of short, stiff, forward directed hairs is golden to orange-yellow colored. In the Guianan white-faced saki P. pithecia the mask is albinotic, white with orange-colored cheeks in males from Guyana and Suriname, and overall white in males from French Guiana. Sakis of the P. monachus and P. pithecia C lades Figure 29. Among the Saki Monkeys genus Pithecia three monophyletic clad is tic Groups or Clades are distinguished: the P. monachus Group containing four taxa ( P. monachus, P. milleri, P. napensis, and P. aequatorialis) , the P. pithecia Group containing three taxa (P. lotichiusi , P. chvySOCephala , and P. pithecia), and the P. llivSUta Group containing four taxa ( P. hirsuta, P. Prorata, P. vanzolinii, and P. albicans) . We only recognize sexual dimorphism as expressed in metachromic characters in the monachus and pithecia Clades (Courtesy of Stephen Nash). On the origin of allopatric primate species 153 distinguish themselves locomotorily from sakis of the third clade - the P hirsuta Clade. A specific locomotor pattern called “vertical leaping and clinging” is performed during foraging and travel- ing in their preferred habitat, which is the discon- tinuous lower canopy and understory of terra firme rain forest. As these sakis have to leap from tree trunk to tree trunk, they are commonly known as “flying monkeys”. In contrast, saki taxa of the P. hirSUta Clade prefer the middle to upper strata of primary rain and seasonally inundated floodplain forests, which strata are interconnected by thick- stemmed vines and hem i-epiphytic climbing shrubs. For that preferred habitat they have adopted a different locomotor pattern, that of horizontal leaping, and quadrupedal running or hopping across thick horizontal branches and boughs. A significant difference in limb proportions between taxa belong- ing to each of the two Clades has been measured, with those of the P. pitheda Group being longer relative to trunk length (H ershkovitz, 1 987a; 1 987b). Another important feature in which the P. hirSUta Clade distinguishes itself from the P. mOYl- achus and P. pithecici Clades is mean group size and sexual dimorphism. Social groups of taxa be- longing to the P. hirSUta C lade are larger and m ulti- male structured, instead of the extended family group that contains only one or sometimes two adult males in taxa belonging to the other Clades. Moreover, contrary to what recent taxonomies Rio Solimoes vdrzea -Parana' do Salsa ' Lago Uauacu L Coarf tern firme Lago Ayapua varzea s' -Rio Pur 6 s Figure 30. Satellite image taken from the region, where the varzea floodplain of the Rio Solimoes borders on that of the Rio Purus. Behind each floodplain are locate d black-water backwater lakes (rias), such as Lago Coari, Lago Uauacu, and Lago Ayapua. A red line indicates where parapatric bald-faced saki Pithecici hlVSUta is encroaching onto huffy saki P. albicans territory. (Below) Portraits of different adult males of Gray’s saki P. hivSUtd. (Above, left) White-masked mutant male P. hirSUta that was seen roaming around alone far in to P. dlbicdHS territory north of Lago Uauacu. (Above, right) Adult male huffy saki P. albicans, note the black face with the showy albinotic eyebrows and white long-haired hood. 154 Marc G.M. van Roosmalen & Tomas van Roosmalen (merely based on museum collections) suggest, we were not able to recognize metachromic sexual dimorphism in any taxon of the P. hirsuta Clade. In the field, we failed to distinguish gender among group members of P. hirsuta, P. irrorata, and P. albicans. N or could we, in captivity, determ ine their gender without up-close examining the saki mon- key’s concealed genitals. Within the Bare-faced Sakis of the P. hirsuta Clade we suggest the least eumelanin bleached overall blackish-gray taxon P. hirsuta to be the nearest to archetypic taxon. It may well have de- rived from a founder-colony of proto -moiiachus that once traversed the Rio Jurua in eastern direc- tion. The following pathways of metachromic blea- ching and allopatric speciation are recognized. From P. hirsuta that occupies the entire interfluve delineated by the Jurua, Solimoes and Madeira Ri- vers, diverged and radiated away in eastern direc- tion taxon P. irrorata after a founder-colony of progressively bleached P hirsuta traversed or cir- cumvented the Rio M adeira (most likely at its upper reaches) during one of the late-P leistocene glacials. Nowadays, taxon P. irrorata occupies the entire interfluve delineated by the Madeira, Amazonas and Tapajos-Juruena Rivers. Its overall coat is advanced eumelanin bleached in comparison with that of P. hirsuta, and albinotic in the distal half of the hood, the hands and feet. Its tail is less bushy, the hairs more curly. Pitheda irrorata has an almost bare face, and its forehead is only halfway covered by an albinotic hood that does not conceal the cheeks and temples. As a result, the monkey’s profile looks more pronounced. Metachromic skin and fur characters of the head that play such an important role in the taxonomy of monkeys like Pitheda, Sapajus and Ateles are often poorly pre- served in museum specimens. Hence, the confusion in most hitherto elaborated taxonomic reviews of these genera. Zoological collections all over the world have lumped m isidentified taxa, such as P. hirsuta and P. irrorata, under the latter. Some lead- ing taxonomists even attribute sexual dimorphism to the B are -faced S akis. From P. hirsuta to the west diverged taxon P. vanzolinii, after a progressively bleached founder-colony of P. hirsuta traversed the Rio Envira. Pitheda Vanzolinii is now confined to the headwaters of the Rio Jurua. It differs in the al- binotic lower limbs and ventral parts that contrast much with the blackish-gray dorsal parts and tail. From P. hirSUta to the north derived the overall near- a lb in otic taxon P. albicans that is pheomelanin bleached orangish-yellow only on the lower limbs. Buffy Sakis P. albicans occupy the northernmost dead-end distribution of the P. hirsUtaClade, which is delineated by the u ntraversable lower Solimoes River in the north, the lower Jurua River in the west, and the lower Purus River in the east. Buffy Sakis are parapatric with the more opportunistic Gray’s Sakis P. hirsuta, from which they once derived. At its southern limit, its distribution shows an open end running across the Rio Tapaua axis. After it tra- versed the Rio Tapaua to the north, Gray’s Saki P. hirsuta was, and still is expanding its range northw ards to the cost of the B uffy Saki P. albicans. This example may well demonstrate that progress- ively bleached to albinotic primate taxa that occupy dead-end distributions will eventually go extinct. East of the Rio Coari and north of the Rio Tapaua - a left-bank tributary of the Rio Purus - we have con- firmed the sympatric occurrence of the taxa P. al- bicans and P. hirsuta, with P hirsuta advancing onto P. albicans as far north as Lago Ayapua (Fig. 30). North of the Ayapua contact zone in territory exclusively occupied by P. albicans, we once spot- ted and photographed a solitary young male, its head pelage resembling that of male White-faced Saki P. pitheda from the Guianas (Fig. 30). We as- sume that this male was a progressively bleached deviant color morph of taxon P. hirsuta that was expelled from or forced to leave its parental group. It may have ventured into adjacent P. albicans territory north of Lago Uauagu. As we have often seen P. hirsuta groups opportunistically penetrating far into white-water floodplain forest (varzea), this metachromic deviant near-albinotic, sexually dim orphic m utant m ale of taxon P. hirsuta in theory could become the founding father of a new taxon. This could happen after this young male would have attracted one or a few P. albicans females to form a small reproductive family group. It then would have to survive making a year-round living in the extensive varzeas found along the south bank of the Rio Solimoes. We have never seen any saki, uakari or other seed-predating monkey occupying that ecological feeding niche in the varzeas that fringe the right bank of the middle Rio SolimSes. Perhaps, this hypothetical scenario may also explain how metachromic sexual dimorphism in primates could have evolved. On the origin of allopatric primate species 155 In figure 3 1, we have visualized the distribu- tions, allopatric speciation, radiation and sup- posedly followed pathways of metachromic bleach- ing in all known Woolly Monkeys, genus LagO- thvix. Woolly monkeys are exclusive matrix terra firm e rainforest dwellers that under normal circum- stances will never enter white-water floodplain forest (varzea). For that reason alone, the distribu- tion of LagOthrix is greatly determined by riverine barriers. Within the genus only one monophyletic Clade is recognized. We consider the saturated eu- melanin, metachromic least bleached Poeppig’s Woolly Monkey taxon La. poeppigii w ith its overall black to dark chestnut-brown coat the nearest to ar- chetypic woolly monkey. In the north, La. poeppi- glV s distribution is confined by the Amazon River, in the east by the Rio Jurua that is also fringed with extensive varzeas, and in the south and west by the foothills of the Andean Mountain range. From La. poeppigii derived in western direction the Peruvian Yellow -tailed Woolly Monkey La. flavicauda, which has (disputedly) been upgraded to its own genus Oreonax. It occurs in parapatry with La. poeppigii , but genetically isolated from it, as it lives in high-altitude Andean cloud forest. With its al- binotic lower half of the circumocular rings, facial muzzle, chin and pheomelanin bleached yellow tail the taxon is following a pheomelanin pathway towards albinotic. From a founder-colony of some- what eumelanin bleached La. poeppigii that tra- versed or circumvented the upper Rio Jurua and then radiated to the east and north, the darkbrown to black headed taxon La. tschudii derived. Its coat is overall dark gray-brown colored, becoming blackish on all five limbs. It occupies the entire interfluve delineated by the Jurua, Solimoes- WOOLLY MONKEYS LAGOTHRIX Lagothrix lagotricha Q Lagothrix poeppigii Lagothrix carta '23 Lagothrix Rio Javan Lagothrix iugens Lagothrix tschudii , Lagothrix Rio Aripuana Lagothrix Rio Jutaf Lagothrix (Oreonax) flavicauda Figure 31. Distributions, allopatric speciation, radiation, and supposed eumelanin pathways of metachromic bleaching in all known Woolly Monkeys genus Lagothrix. 156 Marc G.M. van Roosmalen & Tomas van Roosmalen Amazonas and Madeira Rivers. From La. tschudii in eastern direction diverged the Black-headed or Geoffroy’s Gray Woolly Monkey taxon La. Cana, its entire coat progressively eumelanin bleached, light-gray colored, with a dark-gray to black head. Only as recent as the late- Pleistocene or early Holocene, an advanced eumelanin bleached founder- colony of La. tschudii must have traversed or circumvented the upper Madeira River north of the Rio Ji-Parana (also known as Rio Machado) in eastern direction. It then extended its range by passing the geographic barrier formed by the ex- tensive Tenharim Savanna in Rondonia alongside its southern border. This way, it could enter the interfluve delineated by the Madeira, Amazonas and Tapajos Rivers. Circumventing the extensive Tenharim Savanna, taxon La. Cana apparently missed the narrow entrance to the north that exists between the upper Rio Ji-Parana and the Rio Roosevelt. This could well explain why woolly monkeys are absent from the entire Rios M adeira/A ripuana interfluve north of the Rio M armelos. The relatively recent occupation by La. Cana of the entire interfluve delineated by the Madeira, Aripuana, Amazonas and Tapajos Rivers is near to its completion. Taxon La. Cana’s current northernmost distribution gets to a halt at the lat- itude running across the upper reaches of the Abacaxis and A ndira Rivers, not much south of the untraversable Rio Amazonas. We assume that only when La. Cana invaded all smaller interfluves east of the Rio Aripuana and west of the Rio Tapajos, it began to displace the A 11 -black Woolly Monkey that in the far geological past evolved in the area east of the (proto)-M adeira River. This newly identified woolly monkey still has to be collected and de- scribed. We here provisionally allocate the common name “Rio Aripuana Black Woolly Monkey” to this fully saturated eumelanin, all-black taxon. Ap- parently, as it occupies the same ecological niche as newcomer La. Cana , the Rio Aripuana Black Woolly Monkey finds itself on the verge of ex- tinction. It is smaller, lives in small, socially less complex family groups, and its coat is in meta- chromic respect the most primitive or archetypic. It lives in sympatry with La. Cana, but only hangs on in a small enclave distribution situated between the lower to middle Rio Aripuana and the Rio Acari. It may well represent the ancient, most original, ar- chetypic taxon of all Woolly Monkeys genus LagO- thrix that evolved in the L ate-Pliocene east of the proto-Madeira River, fully isolated from the rest of the Amazon. Woolly monkeys also radiated into the north- western Amazon, most likely after a founder-colony of taxon La. poeppigii circumvented or traversed the upper Amazon River in Peru (where it is called Rio Maranon). Two progressively eumelanin bleached forms that derived from La. poeppigii once must have occupied the Colombian Amazon: the euchromic light-gray Colombian Woolly Mon- key taxon La. lugens that occurs at high altitudes in the foothills of the S outh-C olom bian Andes and in the upper Rio Magdalena valley, and the Brown or Humboldt’s Woolly Monkey taxon La. lagOtricha. The coat of taxon La. lugens is eum elanin bleached charcoal to light-gray colored, but lacks any mix- ture with brown. On the head, a mid-dorsal stripe and a rim across the eyebrows are advanced bleached to euchromic. Mean body size and weight in La. lugens are the largest among all extant woolly monkeys. LagOtvicha' s coat is progressively eu- melanin bleached light-brown colored, except for the blackish hands and feet. Its head is light-brown colored, with a slightly bleached yellowish eyebrow rim and sideburns aside of the blackish-brown face. Taxon La. lagOthricha ranges across the Colom- bian, Venezuelan and NW Brazilian Amazon. Most interestingly, we confirmed the small distribution of a newly identified, advanced pheo- melanin bleached, overall orange-colored taxon in the upper reaches of the Rio Jutai. A founder-colony of advanced pheomelanin bleached La. poeppigii mutants pushed out of La. poeppigii territory must once have successfully adapted to white-water seasonally inundated floodplain forest (varzea) located between the east bank of the upper Rio Jutai and the west bank of the Rio Jurua, near the town of Eirunepe. We were not able to determine the exact range of the Rio Jutai Woolly Monkey, for the area is inhabited by uncontacted Amerindians of the Korubo tribe (so-called “cageteiros”) that are known to kill any non-indigenous intruder. We encountered in the zoological collection of the Brazilian Museu Goeldi (MPEG, Belem - PA ) an overall orange-colored stuffed juvenile specimen that was deposited without collecting data. This very animal is depicted in Da Cruz Lima’s 1945 Mammals of Amazonia. We here provisionally name it the “Rio Jutai Orange Woolly Monkey”. On the origin of allopatric primate species 157 In addition, we found an albinotic overall cream -colored taxon that we provisionally named the “Rio Javan Fair Woolly M onkey” LcigOthrix sp. It resembles much Humboldt’s Woolly Monkey taxon La. lagotricha, but its pelage is longer, softer and silky, besides being overall advanced eu - chromic to cream-white colored. It has long-haired white sideburns alongside a pitch-black face, muzzle and chin. A near-albinotic ancestral fo under- colony must once have been driven out of La. poeppigH territory somewhere near the northern- most border of its distribution. This colony must have been forced to make a living in the white- water floodplain forests (varzeas) that stretch out along the south bank of the Rio Solimoes (near the town of Tabatinga) all the way to the left-bank varzeas of the lower Rio Javari. Under normal cir- cumstances this type of habitat should be conside- red inappropriate for woolly monkeys to guarantee a durable and sustainable living. This seems to be another case where a progressively bleached, near- albinotic founder-colony of La. poeppigH has been driven into a (for woolly monkeys) marginal habitat - seasonally white-water inundated floodplain forest (varzea). According to our theory of allopatric primate speciation, albinotic fair woolly monkeys must have diverged this way from archetypic, sat- urated eumelanin, dark brown coated La. poeppigH. Apparently, it has survived until today in geo- graphic sympatry, but ecological parapatry (inhab- iting adjacent but different habitats) with taxon La. poeppigH , the species it derived from. In 2002, the second author, while at Colombia University, NY, ran the mtDNA sequences of the Rio Javari Fair Woolly Monkey using earlier preserved DNA- samples. He found 4% divergence from sympatric AMAZONIAN SPIDER MONKEYS ATELES Ateles pan isc us Clade 0 A. paniscus Ateles behebuth Clade f A. hybridus A. brunneus 4. beliebuth (Rios NegroIOrinoco) A. variegatus (N PerufSW Colombia] Ateles chamck Clade 0 A. chamek A. sp. nov. (Rios RuruslMadeira) 0 A. longimembris Ewk A. marginatus A. sp, nov. (Upper Rio Xingu) Figure 32. Distributions, allopatric speciation. radiation, and supposed pathways of metachromic bleaching followed in all known Spider Monkeys genus Ateles that occur in the Amazon and along the P acific coast of Ecuador and Colombia. 158 Marc G.M. van Roosmalen & Tomas van Roosmalen Lei. poeppigii and over 7% from the allopatric taxon La. lagOtricha. The AMNH holds three well- preserved skins of the Rio Javan FairWoolly Mon- key LagOtrix sp which were collected by the Olalla Brothers in 1927 along the south bank of the Rio Solimoes, somewhat upstream from the town of Tabatinga. All three specimens are m isidentified as La. lagOtricha (Humboldt, 1 8 1 2 ) . For Spider Monkeys genus AteleS, allopatric speciation, radiation, and phy logeography along different pathways of metachromic bleaching are depicted in figures 32, 33. Four monop hyletic cladistic Groups or Clades are recognized: A. pan- iscus, A. chamek, A. belzebuth, and A. geoffroyi. Spider monkeys have evolved during the Pliocene in the Guayanan Shield, most likely from a pre- cursor of the most ancient of the four extant mono- phyletic cladistic Groups, the A. panisCUS Clade. The Red-faced Black Spider Monkey A. panisCUS from the Guianas represents the nearest to ar- chetypic extant taxon within the genus. This as- sumption is based on some unique primitive characters that are not seen in other spider monkeys. Here we mention: the presence of a vestigial thumb or, if lacking, at least the metacarpal of the first digit that is maintained in the hand; its incapacity of using the tip of the prehensile tail in picking and manipulating small objects like food items; the overall long-haired coat, in particular around the base of the tail and in the forward directed hairtuft on the forehead that resembles a cap; the overall saturated eumelanin black coat without any sign of early eumelanin bleaching; the advanced pheo- melanin bleached bright-red bare face lacking whiskers; the frequent occurrence of albinotic blue- colored eyes; the albinotic cream-white colored, hypertrophied, pendulous clitoris in females and cream-white protruded anus in both sexes, whereas Figure 33. Phylogeographic distribution, allopatric speciation. radiation and metachromic diversification in all known Spider M on key s (AteleS) that occur from the Pacific coast ofW Ecuador and NW Colombia far into C Am erica as far N Mexico. On the origin of allopatric primate species 159 the clitoris is long, flattened and lacking the muscu- lature to erect during foreplay and copulation (Van Roosmalen, 1985a). Mo re over, spider-monkey mat- riarchal social organization is markedly expressed in (leading) female’s body size, which in A. pan- isCUS may exceed that of males; and in the per- manent fu sion -fissio n social structure centered around alpha-females that lead foraging parties on day ranges. As such, complete gatherings of all twenty or so members of a social grouping will never happen (Van Roosmalen, 1985a). This spe- cific type of social organization that is unique am ong N eo tropical prim ates may we 11 be related to the specific phy tosociological composition, pheno- logy and physiognomy of the more ancient, more heterogeneous type of primary terra firme rain forest that evolved uniquely and without major interruptions during the last 60-70 million years on the Guayanan as well as on the Brazilian Shield. Here, available food sources are generally widely dispersed, and rarely clumped at any time of the year. Maturation of nutritious large-seeded fruits - A. panisCUS is a mature-fruit specialist frugivore - is slower and species-specifically stretched out over longer periods of time (Van Roosmalen, 1985b). Mast-fruiting, as commonly seen in tropical rain- forests on other continents, is a phenomenon that does not exist in this ecosystem. Hence, the early evolution of sem i-brachiation (brachiation with the help of a prehensile tail) as the principal locomotor pattern, and the fusion-fission type of social struc- ture during traveling and foraging took place in an- cestral spider monkeys as the principal adaptation of a large-bodied monkey to a well-defined ecolo- gical feeding niche, in a biome that took over 60 million years to develop. It may well explain why the A. panisCUS Clade did not speciate and radiate any further, as the distribution of extant A. panisCUS is still confined to the larger part of the Guayanan Shield. Most plausibly somewhere in the late-Pliocene, from an agouti or saturated eumelanin all-black an- cestor of A. panisCUS derived the phylogenetically distantly related, nearest to archetypic Black Spider Monkey taxon chain ek of the A. chcilfiek Clade. It is distributed south of the Amazon as far south as the Brazilian Shield (in Rondonia and Mato Grosso states). North of the Amazon, the Brown Spider Monkey taxon A. brunneus that ranges in N Colom- bia (in an area confined by the Sierra Nevada Mountains), may represent the least eumelanin bleached, nearest to archetypic taxon of the A. belzebuth Clade. Moreover, in the Pacific coastal forests of Ecuador and Colombia is found the sat- urated eumelanin Brown-headed Spider Monkey taxon A.fusdcepS (form erly A.justiceps filSCWepS). Along the Pacific coast of N Colombia and S Panama is found the all-black but dark red-bellied Colombian Black Sp id er Monkey taxon A. rufiventris (formerly A. fusdceps robustUS). All-black Brown- headed Spider Monkey taxon A. fusdceps may therefore represent the nearest to archetypic taxon of the A. geoffroyi Clade (Fig. 3 2). W ithin the A. chciffiek C lade, nom in ate A. chamek represents the nearest to archetypic taxon. It is sat- urated eumelanin in its overall black coat color and blackish or slightly bleached pinkish circumocular rings and/or facial muzzle, and in the forward directed black hairtuft on the forehead. It ranges across a large part of the Amazon basin delineated by the Amazon River in the north, the Andes Moun- tains in the west, the highlands of the Brazilian Shield in the south, and the Purus and Guapore Rivers in the east. Like the other taxa of the A. chamek Clade, the Black-faced Black Spider Mon- key A. chamek is only found in patches of terra firme rain forest close to major waterbodies, such as lakes, rivers, and creeks. It frequents in particular seasonally inundated marsh forest and black- and clear-water floodplain forest called igapo. We have never spotted spider monkeys belonging to the A. chamek Clade in matrix primary rainforest of the hinterland at distances of over ten km from any major waterbody. There, spider monkeys of the A. chamek Clade are commonly replaced by woolly monkeys ( LagOthvix ) that occupy the same feeding niche in primary terra firme rain forest. All taxa of the A. chamek Clade do laterally migrate to the nearest igapo floodplain forest of clear- and black- water rivers during the 2-3 months lasting fruiting season, which coincides with the peak of the flood. From A. chamek diverged and radiated away in eastern direction the Rio Purus Black Spider Mon- key that we identified to be new to science. This taxon ranges in the interfluve between the Purus and Madeira Rivers, south of the Rio Ipixuna and north of the Rio Tahuamanu in the Bolivian Amazon, a left-bank tributary of the upper Rio Madeira. The Rio Purus Black Spider Monkey AtheleS sp. is having a near-albinotic cream to pink 160 Marc G.M. van Roosmalen & Tomas van Roosmalen colored muzzle, chin, and ears, and a triangular patch of short, backward directed black hairs on the forehead instead of a cap . A f ter a founder- colony of the Rio Purus Black Spider Monkey traversed the Rio Madeira to the east, the Long-limbed Black Spider Monkey A. longimembris diverged. This taxon was already identified as a distinct species by Da Cruz Lima (1945) based on two specimens that were collected by Leo E. M iller along the upper Rio Ji-Parana in M ato Grosso during the first part of the 1914 Roosevelt-R ondon Expedition. It was first de- scribed as Ateles longimembris by Allen ( 1 9 1 4 ) . Holotype and paratype of A. longimembris depos- ited in the zoological collection of the AMNH under No. 36909 were later m isidentified as A. chamek and therefore not included in Kellogg & Goldman’s (1 944) revision of the Spider Monkeys genus Ate- les. The La tin name thatAllen (1914) attributed to this taxon relates to the “ excessively long tail and limbs, the tail length very nearly twice the length of head and body" . A side of its elongated and slender limbs, taxon A. longimembris is further character- ized by the pitch-black face and ears, except for a pale cream-white albinotic triangular patch on the nose, and a wide triangular patch on the forehead that is barely covered with sparse backward direc- ted, stiff, black hairs. Another character of this taxon is the relatively robust incisors and canines that look oversized so that the lips seem unable to conceal them . This feature gives adult Long-lim bed Black Spider Monkeys taxon A. longimembris a bulldog-like appearance. Moreover, its loud or long-distance calls that are so typical for other spider monkeys do not carry far. They sound like bird whistles blowing in the wind. The distribution of A. longimembris is confined by the Rio Madeira in the west, the lower Amazon River in the north, the Rio Tapajos-Juruena in the east and the Rio Ji-Parana in the south. From a founder-colony of A. longimembris that once traversed the Rio Tapajos- Juruena to the east derived the W h ite- w h iskered B lack Spider M onkey A. marginatUS. It is all-black and only euchromic in the small triangular forehead patch or blaze formed by backwards directed white hairs. However, we have seen also adult free-ran- ging A. marginatUS that had black forehead patches. This taxon occupies the interfluve delineated by the Rios Tapajos and Teles-Pires in the west, the lower Amazon River in the north, the Rios Tocantins and Araguaia in the east, and the upper Rio Teles-Pires or Rio M inisuia-M igu (both right-bank tributaries of the upper Rio Tapajos) in the south. After a somewhat eumelanin bleached founder-colony of A. marginatUS once traversed the upper Rio Teles- Pires south of the A. marginatUS distribution, a new taxon diverged that we name the Upper Rio Xingu White-whiskered Brown Spider Monkey. Its coat is chestnut-brown dorsally, and lighter brown on the ventral parts. The snow-white semi-crescent blaze is much larger than in A. marginatUS. It widens above the eyes into long sidewards directed streaks. This newly identified taxon distinguishes itself also from taxon A. marginatUS in the long white whiskers that run from below the eyes across the lips and chin. Moreover, facial skin is pink to flesh- colored in the circumocular rings, muzzle, lips and chin.Within the monophyleticA. chamek C lade the White-whiskered Brown Spider Monkey from the Upper Rio Xingu represents the furthermost eu- melanin bleached taxon that, in accordance with our theory, m etachrom ically and phylogeographically radiated farthest away from archetypic Black-faced Black Spider Monkey taxon A. chamek. W ithin the A. belzebuth Clade, we recognize the dorsally saturated eumelanin darkbrown Brown Spider M onkey taxon A. brunneus as the nearest to archetypic taxon. Belly and inner limbs are eu- melanin bleached light- brown colored. The trian- gular forehead patch formed by backward directed hairs is only slightly bleached brownish-black colored. Taxon A. brunneUS is found in N Colom- bia, between the Cauca and Magdalena Rivers. It is taxonom ically treated as a subspecies of A. hybridus. In the far geological past, the A. belzebuth Clade could well have derived from the archetypic, saturated eumelanin, all-black taxon (A. geoffroyi) A. fusciceps (form eriy A. fusciceps fusciceps ) of the A. geoffroyi Clade that occurs west of the Andes Mountains in the Pacific coastal forests of Ecuador and Colombia. An ancestral founder-colony of A. fusciceps once may have circumvented the Sierra Nevada north of it and diverged into ancestral A. brunneus in the western part of the lower Rio M ag- dalena valley. After a progressively eumelanin bleached founder-colony of A. brunneus traversed the Rio Magdalena to the east, the light-brown and silvery-white colored Variegated Spider Monkey taxon A. hybridus could have derived. It ranges from the northern Colombian Rio Magdalena Basin into the southw esternm ost corner of Venezuela, in On the origin of allopatric primate species 161 the foothills of the Sierra Nevada mountain range (near the city ofMerida). Inner parts of limbs, belly and the small triangular forehead patch are silvery white in taxon A. hybridus , whereas the rest of the coat is light-brown colored. An advanced eu- melanin bleached founder-colony of A. Hybridus once may have circumvented the Sierra Nevada Mountains to the east and reached the headwa- ters of some of the Rio Orinoco’s tributaries in Venezuela’s Amazonas state. It then diverged into the furthermost pheomelanin bleached White- bellied Spider M onkey taxon A. belzebuth. It ranges from north of the Rio Negro and west of the Rio Branco into the Venezuelan State of Amazonas west of the Rio Orinoco, and also far into the lowland Amazon of Colombia. Upper parts, head and dorsal coat of White-bellied Spider Monkeys taxon A. belzebuth are light-brown, but their pelage on ventral parts and inner sides of limbs are silvery white, often pheomelanin bleached yellow to orange-colored. The skin of muzzle and chin is pale brown to pinkish colored. The triangular forehead patch or blaze is light brown, and the eyebrows, whiskers, and throat are silvery. From a founder- colony of A. belzebuth that once traversed the upper Rio Caqueta derived in southwestern direction the southernmost distributed taxon of the A. belzebuth Clade, A. variegatus. This taxon occurs in the N Peruvian, SW Colombian and eastern part of the Ecuadorian Amazon, east of the Andes Mountains and north of the Amazon River (where the river is called Rio Maranon). Its coat is dorsally eumelanin blackish to dark gray, and ventrally euchromic to albinotic silvery-white, except for the dark grayish hands and feet. The legs are silvery white, as are the whiskers and the large blaze or triangular patch on the forehead. Advanced pheomelanin bleached color traits (yellow and orange) as seen in A. belze- buth are lacking in A. VCiriegCltUS. In accordance with our theory and the principle of metachromic bleaching, within the belzebuth Clade the most eu- chromic taxon, A. variegatus, has phylogeograph- ically radiated the farthest away from the dark- brownish colored, nearest to archetypic taxon A. brunneus (Fig. 32). Within the A. geojfroyi Clade (Fig. 33), we re- cognize the saturated eumelanin Colombian Black Spider M onkey taxon A. (jusciceps ) rufiventris (formerly A. fusciceps robustUS ) from the Pacific coastal forests of Colombia and South Panama west of the Andes Mountains as the nearest to archetypic taxon of the A. geoffroyi Clade. Its coat is glossy pitch-black, whereas color morphs of this taxon show a saturated pheomelanin dark red colored belly and genital area. Fur on the fo rehead is slightly brownish tinged. From taxon A. rufiventris derived in southern direction the Brown-headed Black Spider Monkey, the nominate taxon A. (fus - deeps) fusciceps from the Pacific coastal forests of Ecuador and Colombia. It is slightly eumelanin bleached blackish-gray on the belly, brownish black above, with a yellow-brown anterior crown, grading from brown to black on the nape. It often has a white m ustache and beard. Taxa A. fusciceps and A. rufiventris stand at the base of the monophyletic C entral-A m erica S pider-M onkey A. geojfroyi C lade, which radiated away in northwestern direction across the Isthmus of Panama into CentralAm erica as far north as Mexico. From the Colombian Black Spider Monkey A. rufiventris derived the advanced euchromic, near-albinotic (except for the saturated eumelanin feet, hands, lower arms and distal part of the tail) taxon A. ( geoffroyi ) grisescens. However, the validity of this taxon is doubtful, for it has never been seen in the wild. It is thought to occupy a dead-end distribution along the Pacific coast from the Rio Tuyra valley in SE Panama into the Cor- dillera de Baudo in NW Colombia. To the east, its distribution is confined by territory occupied by the Colombian Black Spider Monkey A. rufiventris. From A. rufiventris diverged in western direction the advanced pheomelanin bleached Ornate Spider Monkey taxon A. ( geoffroyi ) panamensis. it is argued that the form A. panamensis is a junior synonym of A. ornatus. Taxon A. panamensis/ OmatUS has a golden brown, dark red to orange colored back, with saturated eumelanin black pe- lage on the top of the head, outer sides of legs, hands, feet and distal part of the tail. It is distributed throughout Panama (from Chiriqui Province as far as E of the Canal Zone) and C+E Costa Rica. From A. ornatus (or A. panamensis ) derived in southern direction the advanced pheomelanin bleached Azuero Spider Monkey taxon A. azuerensis. Its back is gray ish -bro w n , somewhat darker than the underside. Outer surfaces of the limbs are black, the top of the head and neck are (brow nish)-black. Its distribution is delineated by the Panamanian Pacific coast in the south and east. From the Or- nate Spider Monkey taxon A. OmatUS derived in 162 Marc G.M. van Roosmalen & Tomas van Roosmalen northern direction into Nicaragua the advanced pheomelanin bleached, near-euchrom ic Geoffroyi’s Spider Monkey taxon A. geojfwyi. It is silvery to brownish-gray on the back, upper arms, and thighs. Its coat (except for the black head, elbows, knees, upper arms, lower legs, hands and feet) is overall orangish and cream-white colored. Its face is black, often with flesh -colored ‘spectacles’ around the eyes. From A. omatUS radiated away, first in western direction and from coastal Costa Rica northwards into Nicaragua, the advanced pheo- melanin bleached Black-browed Spider Monkey taxon A. ffflntatUS . W ith its orange, black and white coat A. ffflntatUS is the most colorful taxon of the entire A. geoffroyi Clade. From taxon A. frOYltatUS derived the overall most euchromic bleached Mexican Spider Monkey taxon A. VellewSUS. Its dorsal surfaces range from black to light brown, and contrast strongly with its lighter abdomen and inner limbs. Flesh-colored skin is often present around the eyes. It occupies the entire northwestern part of the Isthmus containing El Salvador, Honduras (along the N coast into the lowlands of La Mosquitia), Guatemala (including the highlands) and E & SE Mexico. From taxon A. VelleWSUS to the north derived the near-albinotic Yucatan Spider Monkey taxon A. yucatanensis . It is characterized by the overall advanced eumelanin bleached, light brown and white colored coat. Its fur is brownish- black on the head, neck, and shoulders, grading into lighter brown on the lower back and hips and contrasting with its silvery-white underside, inner limbs, and sideburns. Ateles (geojfwyi) yUCCltan- ensis occupies a large distribution containing NE Guatemala, all ofBelize, and SE Mexico (Yucatan Peninsula). The near-albinotic taxa A. VellewSUS and A. yuCCltanensis that occupy dead-end distribu- tions confined by u ntraversable geographic barriers in the northernmost range of the A. geojfwyi C lade, phenotypically do resemble taxon A. griseSCeilS (from SW Panama) that occupies the southernmost distribution of the A. geoffroyi Clade within the Isthmus. These taxa are equally confined to phylo- geographic dead-end distributions, therefore fully concurring with our theory that pretends to unveil and retrace allopatric primate speciation and radi- ation along phy logeographic pathways of meta- chromic bleaching. Woolly Spider Monkeys or Muriquis genus Brachyteles (family Atelidae) from SE Brazil are disputedly the largest among New World monkeys (adults weighing up to 11-12 kg). It is estimated that alouattines (howling monkeys) and atelines (woolly, spider, and woolly spider monkeys) split about 16 MYA and that the ancestor of Muriquis ( Brachyteles ) and Woolly Monkeys ( Lagothrix ) separated about 10 MYA from the lineage that would eventually lead to the Spider Monkeys ( Ate - les). A m azonian LagOthrix and A tlantic Forest Bra- chyteles are therefore considered to be sister groups (Mittermeier et a 1 . , 2013). Two species ofMuriquis are recognized: the Northern Muriqui B. hypox- anthuS , and the Southern Muriqui B. arachnoides (Fig. 34). Taxon arachnoides is distributed in SE Brazil, through the coastal Serra do Mar in the states of Rio de Janeiro, Sao Paulo, and (the NE of) Parana. Its northern limits are the Serra da Mantiqueira and the Rios Paraiba and Paralba do Sul. Taxon B. hypoxanthus historically ranged through the Atlantic Forest in the states of Bahia, Espirito Santo, Minas Gerais, and Rio de Janeiro, excluding only lowland forests in the extreme S of Bahia and N Espirito Santo. The northern limit of its distribution was probably the Rio Jequiriga or the right bank of the Rio Paraguagu, whereas the southern limit most likely was the Serra da Mantiqueira, in S Minas Gerais state. There, it meets the distribution of the Southern Muriqui taxon B. arachnoides. Sexual dimorphism is absent in Muriquis. The Southern Muriqui has a predom- inantly beige, with light, or dark brown or light gray-brown colored coat. It retains the black pig- mentation of the face, palms, and soles of the feet from infancy into adulthood. Adults of both sexes develop only minor depigmentation in small pink or white spots in the pubic region and sometimes on the face. The Northern Muriqui taxon B. hypOX- anthliS has a uniformly beige colored pelage, with light or dark brown or light gray-brown colora- tion. At birth, the face is black, but at sexual matur- ity face and genitals lose their pigmentation and become spotty pink or flesh-colored (Fig. 34). Northern Muriquis have a vestigial thumb, which character differentiates them from Southern Muriquis that lack the thumb. The Southern M uriqui seems to be nearer to archetypic woolly spider mon- keys than the Northern Muriqui, for the latter is overall progressively pheomelanin bleached near- albinotic in the head characters (white eyebrows, sideburns, and beard), and also in the advanced de- On the origin of allopatric primate species 163 Brachyieles hypoxanxhus Brachyreles arachnoides - 30 ' Figure 34. Metachromic bleaching in the Woolly Spider Monkey or Muriqui genus Brachyteles from the Atlantic coast of SE Brazil. pigmentation of the face, in particular the spotty flesh- colored muzzle (Mittermeier et al., 2013). For Amazonian Howling Monkeys, genus Al- OUCltta, allopatric speciation, radiation, and phylo- geography along eumelanin and pheomelanin path ways ofmetachromic bleaching are depicted in figures 35, 36). Two monophyletic cladistic Groups or Clades are recognized: AL belzebul and AL seniculus (M itterm eier et al., 2013). W ithin the Al. belzebul Clade, distributed south of the Amazon, saturated eumelanin all-black howling monkeys of the Amazon Black Howler taxon Al. nigerrima range between the Tapajos and Madeira Rivers. A founder-pair or colony of somewhat bleached Al. nigerrima howlers once must have traversed the lower Rio Madeira, most likely lifting on floating logs or on drifting islands covered with chavascal (low type of varzea) forest. Presently, this howler also inhabits almost the entire interfluve delineated by the Rios Amazonas, Purus and Ipixuna, an area that was formerly occupied by the advanced pheo- melanin bleached yellowish-orange colored Purus Red Howler taxon AL pUTUensls (belonging to the AL seniculus Clade). We have spotted Al. nigerrima howlers in the varzea near Carreiro (opposite the city of Manaus) and, also, as far south as the Rios 164 Marc G.M. van Roosmalen & Tomas van Roosmalen Igapo-Agu and Tupana - black-water rivers that empty out into the Rio Madeirinha (a white-water left-bank tributary of the Rio Madeira). It seems that the overall orange-colored resident howler Al. puruensis and the all-black invasive Al. nigerrima howler do co-exist locally. However, the two taxa do not mix nor interbreed. While conducting a canoe survey during the peak of the flood season in the vast igapo floodplain along the Rio Igapo-ACI:ide A to not to seme it I us (lade sum sub- f t ii do IQ inches mgettma discotor uHiwo male uhilata Fetnale Atouattn bet zebu l Clade Figure 36. Metachromic variation, radiation, and phy logeography along eumelanin and pheomelanin pathways of metachromic bleaching depicted for all known Amazonian Howlers of the Alouatta. SeniculuS and Al. belzeblll C lades . 166 Marc G.M. van Roosmalen & Tomas van Roosmalen Figure 37.Adultmale Guianan Red Howler AlouattCl mac- connelli. It was photographed while pulling off a juvenile that tried to seek protection from the human intruder, some- where along the black-water Rio Jauaperi, a north-bank tributary of the Rio N egro (Courtesy of D avid Lem m on ). red coat and a rufous-chestnut dorsal band, hands, feet, and tip of the tail. A further pheomelanin bleached founder- colony of Al. discolov must once have traversed its eastern distributional limit - the Rio Xingu, Iriri, or Santa Helena (left-bank tributary of the Rio Teles-Pires in Mato Grosso). From Spix’s Howler Al. disColOT derived the dark brown colored Red-handed Howler Al. belzebul. This species is progressively pheomelanin bleached in the reddish-brown to yellow hands, feet, tail tip, forehead and back. It is distributed south of the Amazon, east of the Rio Xingu-Iriri, in the states of Para (including Mexiana, Caviana, and M arajo Islands in the Amazon estuary), Maranhao, Tocantins, and Mato Grosso. West of the Atlantic coast of NE Brazil in the states of Rio Grande do Norte, Paraiba, Pernambuco, and Alagoas, are found enclave populations isolated from what is thought the taxon’s former distribution, which must have been continuous through the states of Ceara and Piaui to the Amazonian population. From the Red-handed Howler Al. belzebul derived the ad- vanced pheomelanin bleached Maranhao Red- handed Howler Al. ululdtd. This species distributed in NE Brazil occurs in remnant forest patches of dry forest scrub called caatinga. It enters also the coastal mangrove forests of northern Maranhao, Piaui, and Ceara. The Maranhao Red-handed H ow ler Al. ululdta. radiated farthest away from the archetypic overall black Central Amazon Black Howler Al. YligewilYlCl. It is sexually dichromatic. The male is black with rufous to reddish-brown hands, feet, tip of the tail and flanks. The female is yellowish-brown with sparse grayish hairs, giving it an overall olivaceous appearance (Fig. 36). Within the Al. seniculus Clade we consider the overall dark reddish-brown Ursine Red Howler arctoidea from N Venezuela east of Lake Mara- caibo, and from the coast (including the Islands of Trinidad and Tobago) extending S thro ugh the llanos to the Rio Orinoco, the nearest to archetypic taxon for the m onophyletic Al. seniculus sub -Clade, which is distributed north of the Amazon. Both sexes have a coat that is dark reddish-brown on the body, contrasting with a darker brown to blackish head, shoulders, limbs, and proximal part of the tail. Male Ursine Red Howlers often have a blackish beard, limbs and tail. From a founder-colony that once traversed the Orinoco River to the east, has de- rived the advanced pheomelanin bleached Guianan Red Howler Al. ITICICCOnYielli. This taxon ranges east of the Rio Orinoco throughout the Guianas, N Brazil (east of the Rio Negro and north of the Rio Amazonas, including Guru pa Island in the Amazon estuary), and S Venezuela (between the Cassiquiare and Orinoco Rivers). The Guianan Red Howler’s coat is uniformly dark rufous-brown, the back is pheomelanin bleached yellowish to golden-brown with a dark dorsal stripe, and arms to elbows and legs to thighs are orangish-red . Distal part of the tail is pale-yellow (Figs. 35, 36). From a founder-colony of further pheomelanin bleached Al. macconnelli that once traversed either the upper Rio Negro in the Colombian Amazon, or the Orinoco River at its headwaters, has derived the overall orange-colored Colombian Red Howler Al. Seniculus. This taxon is now distributed north of the Amazon across E Ecuador and E Peru (east of the Rio Huallaga), Colombia, NW Venezuela, and the Brazilian Amazon inbetween the Rio Solimoes and Rio Negro. The Colombian Red Howler Al. Sen- iculus is overall golden-toned to coppery-red on the body, contrasting with the maroon head, shoulders, lim bs, and proxim al part of the tail. M ale Colombian Red Howlers are much bigger than fe- On the origin of allopatric primate species 167 males. Within the monophyletic Al. SBTliculllS sub- Clade the bright orange-red Colombian Red Howler Al. seniculus is phylogeograpically the most ad- vanced pheomelanin bleached taxon. It radiated the farthest away from the nearest to archetypic satur- ated eumelanin, overall dark brown colored Ursine Red Howler Al. ClfCtoideci, as such fully concurring with our theory. With respect to the monophyletic Al. Sara sub-Clade of the Al. SeiliculliS cladistic Group that is largely distributed south of the Amazon, we consider the Jurua Red Howler juara the nearest to archetypic, least eumelanin bleached (dark brown) taxon. It ranges in the W Brazilian Amazon south of the Rio Solimdes, and in the Rio Jurua Basin, extending west into the Peruvian Amazon. It is not sexually dichromatic. Its coat is generally dark reddish-brown, with the middle of the back lighter orange-rufous colored, and limbs and tail base dark rufous to black. The tail is paler, more golden from middle to tip. From a pheo- melanin bleached founder- colony that once tra- versed the Rio Jurua to the east derived the Purus Red Howler taxon Al. purueiisis. It is distributed across the entire Rios Jurua/Purus interfluve as far east as the middle Rio Madeira. From there, it ex- tended its range across the upper Rio Aripuana as far east as the Rio Teles-Pires, and south as far as the Rio Abuna (which forms Bolivia’s northern border). The Purus Red Howler Al. puruensis is sexually dichromatic. Males are dark rufous or red- brown with a golden upper dorsum and shoulders, whereas females are golden-orange with distal por- tions of limbs, tail base, and beard dark rufous. From a progressively pheomelanin bleached founder-colony of the Purus Red Howler that once traversed the Rio Abuna, has derived the quite distinct Bolivian Red Howler taxon sara. It is dis- tributed across the Bolivian Amazon including the entire Rio Beni Basin, and east as far as the Rios M am ore/G uapore interfluve. The Bolivian Red Howler’s coat is brick-red above, with limbs, head, and proximal part of the tail darker, more rufous colored. It represents the most advanced pheo- melanin bleached taxon of the Al. Sara sub-Clade (Fig. 36). It occupies a dead-end distribution in the south bordering the drier savanna and Chaco area (Fig. 35). Going further southwards begins the dis- tribution of the Paraguayan Howler Al. caraya. For extra-A m azonian Howling Monkeys genus Alouatta, allopatric speciation, radiation, and phylo- geography along eumelanin and pheomelanin pathways of metachromic bleaching are depicted in figures 38, 39. Four non-Am azonian monophyletic Clades are recognized: the Brazilian Brown Howler Al. guariba, the Paraguayan Howler Al. caraya , the C entral A m eric an M an tied H o w ler Al. palliata, and the Mexican Black Howler Al. pigra (M it term eier et al., 2013). The Brown Howler Al. guariba C lade consists of two populations that may represent dif- ferent valid taxa or species: the Northern Brown Howler Al. guariba and the Southern Brown Howler clamitans. Taxon Al. guariba ranges in the Atlantic Forest from the Rio Paraguagu, Bahia State, along the coast south as far as Rio Paraiba in Rio de Janeiro State. Inland, it extends into Minas Gerais State. The Southern Brown Howler Al. claiTl- itans is distributed in the Atlantic Forest south of Rios Doce and Jequitinhonha, south as far as Rio Grande do Sul State. Taxon Al. guariba is not sexu- ally dichromatic and both sexes are red-fawn, with females usually somewhat duller in color. Taxon Al. clamitans is generally dark reddish-brown, with males often being lighter colored than females. Males from Sao Paulo are orange-red to red-brown with a red belly, whereas males from Santa Catarina and Rio Grande do Sul are bright red-orange, having dark brown feet. Females are overall dark brown or blackish. The Northern Brown Howler Al. guariba is the lesser metachromic bleached. Taxon Al. clamitans derived from it, the males progress- ively following the pathway of pheomelanin bleaching. The further south it ranges, the more the male’s overall coat color tends to red-orange or bright orange (Fig. 39). The Paraguayan Howler Al. Caraya forms a mono ty pic Clade. It is a sister species to the Am azo- nian red howlers of the Al. seniculus Clade. It di- verged from a common ancestor about 4 MYA. The Paraguayan Howler Al. Caraya is distributed across C Brazil, south of the states of Para, Tocantins, M aranhao, and Piaui, west into the Pantanal, south into Paraguay, E and SE Bolivia, and maybe also into NW Uruguay. Much of its range is in the ‘cerrado’ of central Brazil and semi-arid ‘caatinga’ forest scrub in NE Brazil, where it uses gallery and riparian forest and patches of seasonal (semi)- deciduous ‘cerradao’ (a type of savanna forest). Adults of Al. Caraya are sexually dichromatic, but both sexes are blond at birth. Mature males are generally uniformly black. Females and young of 168 Marc G.M. van Roosmalen & Tomas van Roosmalen either sex are pale grayish-yellow to golden-brown. Male Paraguayan Howlers AL CCITCiyCl from Bahia and Goias are black, but those from Mato Grosso and Parana are black with a brown back and hind parts. Males from Sao Paulo and Minas Gerais States are brown-black, with yellowish hands, feet, belly, and tail tip. In all male individuals the face is invariably dark, the fur is stiff and lengthy, and the beard is prominent. The scrotum is rust-red colored. The Pacific Coastal and Central American Mantled Howler Al. pCll licit Cl is, based on geo- graphic distribution, divided in five taxa that could well represent distinct valid species: AL palliata from NE Guatemala, ranging east to E Costa Rica or W Panama; Al. ClCCJUCltonCllis from the southern distributional lim its of Al. palliata ranging through the D arien into W Colombia, W Ecuador, and south as far as NW Peru; Al. mexicCUia ranging from S to SE Mexico and Guatemala following the southern- most distribution of the Central American Black Howler Al. pigva\ Al. COibensis from Coiba and Jicaron Islands in SW Panama; and AL trabeata from the Azuero Peninsula in SW Panama (Fig. 38). The coat of the Mantled Howler is smooth, very short and upright, being silky black with a mantle of longer, gold or yellowish-brown fur along the flanks. Adult males have a white scrotum. The Central American Black Howler AL pigTCl is monotypic. It is distributed across SE Mexico, Belize, and N to C Guatemala. Fur of AL pigTCl is notably long, soft, and dense. Adults are not sexu- ally dichromatic. They are overall black with traces of brown on the shoulders, cheeks, and back. The EXTRA AMAZONIAN HOWLERS ALOUATTA Ala it alia pigra Clatle < Alouatta pigra Alouatta palliata Clade Alouatta (p.J mexicana Alouatta (p.) palliata Alouatta (p.) aequatorialis Alouatta (p.) trabeata Alouatta (p i eoihemh Alouatta caraya CJade Alouatta caraya Alouatta guariba Clad c jt — Q Alouatta (g.) guariba 'f Alouatta (g.) clamitam A jJifTc ,4. p- acbtctu A p pillmtc A etrtfe A f. A l tUrnimw Figure 38. Distributions, allopatric speciation, radiation, and pathways of metachromic bleaching followed in all extra- Amazon ian Howling Monkeys genus AloUClttCl. Two Clades occur south of the Amazon: Al. gUCiribci from the E Brazilian A tlan tic forest, and Al. CCLTUyCl from the ‘cerrado ’ and ‘cerradao ’ of the Central Brazilian Plateau. Along the Pacific coast of Ecuador and Colombia, far into Central America, occur the Al. pcillicitci and Al. pigra Clades. On the origin of allopatric primate species 169 Figure 39. Pelage color variation, radiation and metachromic bleaching along eum elan in and pheomelanin pathways depicted for all extra- A m azo nian Howling Monkeys genus AloilClttCl of the Brazilian Al. guaviba, and Al. CCIVCiyCl C lades, and the C entral-A m erican Al. pigra and Al. palliata Clades. 170 Marc G.M. van Roosmalen & Tomas van Roosmalen CentralAmerican Black Howler Al. pigra is consi- dered the most saturated eumelanin, least bleached, nearest to archetypic form of the Al. pigra and Al. palliata Clades. It is also by far the largest how ling m onkey. The Al. palliata C lade is believed to have diverged from ancestral Al. pigra about 3 MYA (Mittermeier et al., 2013). Taxon Al. palliata is sympatric with taxon Al. pigra in Tabasco State, Mexico and in a small part of Guatemala. From the Golden-mantled Howler Al. palliata radiating northwards derived the Mexican Howler A/, mexi- Cana , and radiating southwards the South Pacific Blackish Howler Al. aequatorialis, ranging far south into the Pacific coastal forests of Colombia and Ecuador. There, it is sympatric with the Colom- bian Red Howler Al. seniculus. From Al. aequatO- rialis in SW Panama derived the Azuero Peninsula Howler Al. trabeata and the Coiba Island Howler Al. coibensis (Fig. 3 8). During his long-term fieldwork on the ecology of all eight monkey species that occur in the Guianas, the senior author has repeatedly watched the basic principles of allopatric primate speciation atwork.Athis study site situated in pristine prim ary terra firm e rain forest in Central Suriname, local populations of the Guianan Red Howler Al. UiaC- connelli (Fig. 37), the most territorial among all extant howling monkeys when measured by the size of the hyoid bone, had passed beyond the howler’s optimal densities (Van Roosmalen, 2013a; 2015). This was measured by the high frequency of dawn chorus and vocal battles of neighboring groups throughout the day and nighttime in the proximity of territorial boundaries. One day, a subadult male got pushed out of his parental group that ranged close to the campsite area. For several days after being expelled, this young howler male got repe- atedly involved in vocal battles with neighboring groups that subsequently chased him out of their respective territories. Weeks later, far away from the campsite, a boundary conflict took place that seemed never ending. The researcher rushed over to the spot. He arrived just in time to witness this very subadult male being attacked by the leader of a resident group in the company of his harem. The whole group chased the young male into an isolated tree close to where he could watch the scene. The subadult male was in the company of a female he presumably had attracted (‘stolen’) from some re- sident group that had chased him out earlier. In an attempt to escape from his attackers, the howler male almost fell out of the canopy. He just could get hold on a thick branch and was hanging under- neath it only secured by the grip of his hands and tail. Then, they all began to bite in his hands and tail tip. With a scream, he let loose and came crashing over forty meters down to the forest floor, hitting it at a hair width away from the researcher’s head. The monkey looked dead, his motionless body covered with blood. After a few minutes, however, he got back on his feet and slowly clim- bed up a small tree. Back in the canopy, he sat next to his mate that had been watching the show from a distance. The pair was never seen again within the borders of the 400-ha study area. Some time later, vocal battling recommenced. It came from the same direction, sounding only much farther away. In retrospect, we assume that the couple survived and in the long run found a place to settle down, start a family, and defend a small territory squeezed inbe- tween the territories of some resident howler groups far away from their respective parental groups. One may speculate that the howler pair, driven by the trend to allopatry, also may have survived by ven- turing into some ‘empty’, marginal, or for howlers unfamiliar habitat. Or, in case the male was expel- led from his parental group for his skin or (part of) coat color being somewhat lighter, he could have joined other outcast males that were discriminated upon and pushed out of their parental groups for other mutant metachromic deviances of skin and/or coat characters. For the sake of survival alone, such healthy young individuals may join efforts to stay alive. Together, they may turn into potential foun- der-colonies venturing into new lands, where they can thrive and reproduce unrestrictedly. At least as long as those lands, in turn, do not reach the taxon’s optimal population density. By the time they do so, the generally accepted phenotype of that new para- patric or allopatric taxon or (eco)-species will have been stabilized while showing whatever features of further metachromic bleaching and/or depilation. Fiving on an island in the Coppename River at about ten km from his field site, the senior author has repeatedly witnessed the coming and going of small groups of potential howler-founders to and fro Foengoe Island after having been forcefully pu- shed out from some mainland territory by the ruling group male(s). Pushed against the riverbank, they apparently did overcome their natural fear of water On the origin of allopatric primate species 171 and then swam toward the island. For some time, such immigrants tried to make a living on the island. Until it became clear to them they were trapped on an island too small to sustain a howler group year-round. Occasionally, such groups were spotted later while ranging along the opposite river- bank. We assume they had traversed the river swim- ming. Interestingly, a female howler that was raised as a pet and then set free to range across the 30-ha island, was eager to join any howlers coming onto the island. Sadly, when the immigrants eventually swam back to the mainland in search of new lands, the female stayed back on the island. Perhaps, she did so for lack of sufficient bonding or for fear of swimming across the river. Capuchin Monkeys (genera CebllS and Sapajus ) have diverged from Squirrel Monkeys (Saimiri) about 15 M YA. They form ed distinct monophyletic Clades that diverged during the Late Miocene to Early Pliocene, about 6.2 MYA. The Clades diver- sified during the Plio-Pleistocene era into two groups: Gracile or Untufted Capuchins (genus CebliS ) in what is today the western Amazon, about 2.1 MYA, and Robust or Tufted Capuchins (genus SapajllS ) in what are today SE Brazil, E Paraguay, and N Argentina, beginning about 2.7 MYA (M it- termeier et al., 2013). Gracile Capuchins genus CebllS are separated into the following five cladi- stics Groups or Clades: Humboldt’s White-fronted Capuchin Ce. albifrons with four+ taxa ( Ce . albi- frons, Ce. yuracus, Ce. unicolor, and Ce. cuscinus ), G uianan Weeper C apuchin Ce. olivaceus w ith three taxa (Ce. brunneus, Ce. olivaceus, and Ce. casta- neuS), White-faced Capuchin capucinus with two taxa (Colombian White-faced Capuchin Ce. Cdpu- dnus and Panamanian White-faced Capuchin Ce. GRACILE CAPUCHINS CEB VS Cebus capucinus Clade Cebus imitator • Cebus capucinus Cebus albifrons Clade • Cebus albifrons 0 Cebus yuracus • Cebus unicolor Cebus cuscittus Cebus aequatorialis Cebus versicolor CLade 0Cebus malitiosus Cebus cesarae Cebus versicolor 0 Cebus leucocephalus Cebus olivaceus Clade Cebus brunneus Cebus olivaceus 0Cebu s castaneus lU/TmiClX Cebus kaapori ( V \ Figure 40. D is tribu tio ns, allopatric speciation, radiation, and pa th ways of metachromic bleaching in all hitherto recognized Gracile Capuchins of the five distinguished phy logeographic Clades: CebllS CCLpUCiflUS, C. olivaCCUS, C. Versicolor, and C. albifrons. The fifth Clade C. aequatoriaUs is monotypic. 172 Marc G.M. van Roosmalen & Tomas van Roosmalen GK WILE CAPl CHINS GEM'S CEBL S 10 inches matitiosus cesaree leucocophaUis Figure 41. Metachromic diversification along eumelanin and pheomelanin pathways of metachromic bleaching (arrowed lines), speciation and radiation in all hitherto recognized Gracile Capuchins (CebuS) of the five distinguished phylogeo- graphic c lades: Cebus olivaceus, C. versicolor, Cebus capucinus, and C. albifrons. Cebus aequatorialis from w Ecuador and NW Peru is monotypic, but may have derived from ancestral C. yUVCLCUS that once traversed the Andes Mts. On the origin of allopatric primate species 173 imitator), and Varied W hite- fronted Capuchin Ce. versicolor with four taxa (Ce. versicolor, Ce. leu- cocephalus, Ce. cesarae, and Ce. malitiosus). The recently discovered Ka’apor Capuchin Ce. kaapori ranging S of the lower Rio Amazonas is geo- graphically closest related to the Guianan Weeper Capuchin (i.e., taxon Ce. COStaneuS ranging along the left/north bank of the lower Rio Amazonas) and, therefore, may form a sister Clade to it (Figs. 40, 41). Within the Ce. capucinus Clade we consider the Colombian White-faced taxon Ce. capucinus (ranging from E Panama, through W Colombia south as far as NW Ecuador), the nearest to arche- typic, least metachromic bleached form. Its body, crown, limbs, and tail are black. The chest is white, extending forward to the face and front of the crown and upward to the shoulders and upper arms. The Gorgona White-faced Capuchin Ce. CUTtUS CUrtUS is a small and relatively short-tailed subspecies from Gorgona Island sitting on the Colombian Pacific coast. From taxon Ce. Capucinus derived Ce. imitator, the taxon that ranges from N Hon- duras, C and W Nicaragua, Costa Rica south into W Panama. It resembles much the typical Colombian White-faced Capuchin Ce. CapudnuS, but females have elongated frontal tufts with a brow nish tinge. Within the Ce. olivaceUS Clade we consider the Venezuelan Brown Capuchin Ce. brumWUS fro m N Venezuela east of the Sierra de Perija and along the Coastal Range, including the island of Trinidad (where it is possibly introduced), the nearest to ar- chetypic, least metachromic bleached form. Its pel- age is thick and long, the upperparts are generally darker along the middle of the back than on the sides, the hairs are dusky basally, with a broad zone of chestnut in the middle, and black at the tips. Face and sides of the head are pale yellowish gray. The crown has a broad V-shaped patch of long hairs, narrowing to a point in front of which a narrow black line runs forward to the nose. Chin and lower parts of cheeks are grayish or fulvous white to whitish. Underparts are blackish brown, with tips of the hairs hazel. The throat is lighter than the chest and belly. Upper arms are maize yellow. Outer forearms are blackish with yellowish tips, inside forearms are much darker. Hands are blackish, hindfeet are nearly black. Tail is colored as back. From ancestral Ce. brunneUS derived the Guianan W eeper C apuchin Ce. olivaceUS that is restricted to the Venezuelan Amazon Basin in forests of the Gu- ayanan Shield, from the upper Rio Orinoco east to the left bank of the Rio Essequibo in W Guyana. Its pelage is overall dark brown or reddish with black- agouti banding on flanks, limbs, and tail. The face is naked and pink. Cheeks are buffy-white. It differs from Ce. brunneUS in the advanced bleached alb i- notic head and upper arms and the wider V-shaped black crow n cap . From ancestral Ce. olivaceUS derived the Chest- nut Weeper Capuchin Ce. COStaneuS. This taxon ranges from the Rio Essequibo E through Suriname and French Guiana into N Brazil, where its distri- bution is delineated by the Rios Negro, Branco, and Catrimani in the W, Rio Amazonas in the S, and the Atlantic coast in the E (it also inhabits Caviana and Mexiana Islands in Amazon’s estuary). It differs from Ce. olivaceUS in the narrower black triangle on the crown and the pelage of the head being ove- rall yellowish-white, but reddish-chestnut above the ear and nape, in the advanced pheomelanin ble- ached reddish-chestnut upperparts of the body and limbs, and pale yellow shoulders and fronts of arms above the elbows.A founder-colony of the Chestnut Weeper Capuchin Ce. ( olivaceUS ) COStaneuS must once have traversed the lower Rio Amazonas. From it derived the Ka’apor Capuchin Ce. kaapori that ranges in NE Brazil south of the lower Amazon River (NE Para and NW Maranhao). This taxon is characterized by a longer body in comparison to other Cebus species. It is grayish agouti-brown, and lighter on the flanks. Face, shoulders, mantle, and tail tip are silvery-gray, the limbs are agouti, and the hands and feet dark brown or black. The crown has a triangular black cap that extends to a dark stripe down the nose. The pelage of the Ka’apor Capuchin is overall advanced eumelanin bleached to nearly albinotic, as such much contrasting with the satu- rated eumelanin blackish crown cap, hands and feet. Being phylogeographically farthest radiated away from the center of dispersion (NW Venezuela) of the Ce. olivaceUS Clade, and occupying a dead-end distribution, where it also has to compete with the Guianan Brown Capuchin Sapajus apella (Figs. 42-44), Ce. kaapori is clearly the most progressi- vely bleached, near-albinotic taxon within the Ce. olivaceUS Clade (Fig. 41). W ithin the Ce. Versicolor C lade w e consider the Varied White-fronted Capuchin Ce. Versicolor the 174 Marc G.M. van Roosmalen & Tomas van Roosmalen ALL ROBUST CAPUCHIN MONKEYS SAPAJUS Sapajus ape l In ( lade # Sapajus a pel la O Sapajus apella margaritae Sapajus macrocephalus Clade Sapajus macrocephalus 4 ssp, Sapajus nigritus Clade # Sapajus nigritus + cucullatus Sapajus cay # Sapajus libidi nosns Sapajus robust us Sapajus xa n thostern os Sapajus flavins Figure 42. Phylogeography, allopatric speciation, radiation, and pathways of metachromic bleaching followed in all hitherto recognized Robust (Tufted) Capuchins of the three distinguished phylogenetic Clades: Sapajus IligritUS, S. apella , and S. macrocephalus. From an ancestral saturated eumelanin (all-black) form of the S. nigritUS Clade , quite recently (an estimated 400,000 YA) radiated away into the Amazon the pheo melanin bleached species S. apella (including the insular taxon S. a. margaritae ), and S. macrocephalus , the latter with four different taxa/’species-in-the-m aking' - from SE to N: juruaUUS, pallidus, maranonis, and fatuellus). nearest to archetypic, least metachromic bleached taxon. It is distributed in N Colombia in the middle Rio Magdalena Basin. It is the darkest among the Clade’s four taxa, though a rather pale form with red tones on the mid-dorsal region and foreparts of the limbs, generally contrasting with the rest of the body (Fig. 4 1). From Ce. Versicolor derived towards the NE the Sierra de Perija White-fronted Capuchin Ce. leUCOCephaluS that ranges in N Co- lombia from the W slope of the Cordillera Oriental E to the Rios Zulia and Catatumbo Basins and NW Venezuela (Zulia State). This taxon is progressively bleached, near-albinotic in the head, chest, and shoulder parts. From Ce. Versicolor derived towards the N first the Rio Cesar White-fronted Capuchin Ce. cesarae, ranging in N Colombia, in the Rio Cesar Valley, W into the S and E slopes of the Sierra Nevada de Santa Marta. From taxon Ce. cesarae, On the origin of allopatric primate species 175 ROBUST CAPUCHINS SAPAJUS Figure 43. Radiation and metachromic diversification in the Sapajus nigritUS and S. Cipellci Clades of Robust or Tufted Capuchins. From an ancestral saturated eumelanin form of S. nigritUS the species Sapajus apella and S. maCWCephalllS radiated away into the Amazon with different taxa ‘in -the-m aking ’ . 176 Marc G.M. van Roosmalen & Tomas van Roosmalen in turn, derived the Santa Marta White-fronted Capuchin Ce. lYialitioSUS that is only known from the NW base of Sierra de Santa Marta. It may range also throughout the lower W and N slopes of the Sierra Nevada in N Colombia. The two taxa are the palest among the N Colombian and Venezuelan White-fronted Capuchins. Taxon Ce. cesarae is buffy in the head and throat parts and pheomelanin bleached orangish in the cap, middle of the back, forearms, and forelegs, as such contrasting with the sides of back and trunk. Taxon Ce. malitioSUS is advanced eumelanin bleached in the silvery to cin- namon-brown chest and belly, and a contrasting al- binotic area of the front extending well over the upper surfaces of the shoulders and inner sides of upper arms (Fig. 41). Within the Ce. albifrons Clad e we consider the M aranon W hite-fronted Capuchin Ce. yurCICUS the nearest to archetypic, overall least metachromic bleached taxon (Fig. 41). This taxon is distributed north of the Amazon River in S Colombia, E Ecuador, NE Peru, and presumably W Brazil between the Rios Iga and Solimoes. It is gray- fronted on the forehead, sides of the face, chest, and outer sides of the arms. Its general color is ochreous-brow n, contrasting sharply with the gray- ish to buffy outer sides of forelimbs, and with the pale silvery to orangish underparts. The cap is black, with a median line running down inbetween the eyes. The tail is brown like the back, but paler tow ards the tip. From ancestral Ce. yUTCICUS derived first Humboldt’s W hite-fronted Capuchin, the nom- inate taxon Ce. albifrons that is widely distributed across the upper Amazon Basin of S Venezuela, S and E Colombia (occurring north of the Rio Amazo- nas and the Rio Iga-Putum ayo, N as far as the Rio Meta, and in the lowlands W of the Orinoco, and NW Brazil (N of the Rio Solimoes, and W of the Rios Negro and Branco, as far north as the Rio Ur- aricoeira). Hum bold t’s W hite-fronted Capuchin Ce. albifrons is overall pale grayish-brown, darker on the limbs. Hands and feet are yellowish-brown. The tail is ashy above, whitish below, and brownish- black towards the tip. The front is creamy white, and there is a cap of short dark fur on the crown that is rounded in the front and well demarcated from the light-colored forehead. The face is naked and pinkish, flesh-colored. From Ce. yuraCUS derived also Spix’s White- fronted Capuchin Ce. Unicolor, most likely after a founder-colony of ancestral Ce. yuraCUS traversed the upper reaches of the Rio Ucayali in E Peru. It is nowadays widely distributed in the upper Brazilian Amazon Basin, south of the A m azon River and w est of the Rio Tapajos, through- out the northern parts of Mato Grosso and Rondo- nia States, and throughout the Rios M adeira, Purus, Jurua, and Javari Basins as far west as the Rio Ucayali. CebliS Ullicolor is uniformly bright ochreous or grayish-brown with darker grayish- brown flanks and mid-back, with a yellowish or cream-white front and reddish-yellow to reddish limbs and tail. From ancestral Ce. linicolor deriv ed later the Shock-headed Capuchin Ce. CUSCiniiS that is believed to range from the right bank of the upper reaches of the Rio Purus in SE Peru, W into the Cuzco Department including the upper Rio M adre de Dios, and S and E as far as the Rio Tambopata Basin, also extending into NW Bolivia. Taxon Ce. CUSCinus has a longer, silkier fur than Ce. linicolor and is less brightly colored. Its limbs are browner and contrast less with the back. The cap is large, distinct, and dark brown. The forearms are orange- rufous on the outside, darker on the wrists and hands. Underparts are ochreous- orange and silvery, becoming buff on the chest. The fronts of the shoulders and inner sides of the upper arms are whitish. The tail is brown, somewhat paler towards the tip. The male has a broad pale frontal region sharply defining the dark-brown cap. Overall, Ce. CUSCiniiS is the most advanced pheomelanin bleached taxon of the Ce. albifrons Clade. It is the form that radiated away farthest from the center of this Clade’s dispersion (Fig. 40). The Ecuadorian White-fronted Capuchin Ce. aequatorialis is mono- typic. It may form a sister Clade to the Gracile Capuchins from the upper Amazon B asin . A ncestral M aranon W hite-fronted Capuchin Ce. yuraCUS once must have traversed the Andes Mountains some- where at the upper reaches of the Rio M aranon and then diverged into Ce. aequatorialis. Cebus aequat- orialis is distributed in Ecuador and NW Peru, in the lowlands west of the Andes (Fig. 34). Its upper- parts are pale cinnamon rufous, darker along the midline of the back. Front and sides of the head are yellowish white, with a narrow black transverse line on the forehead forming the cap, from which a narrow median black line descends to the nose. Hands and feet are a little darker, more brownish than the arms and legs. The chest is lighter than the belly (Fig. 41). On the origin of allopatric primate species 111 During long-term fieldwork in Central Suri- name, the first author spotted a few times by chance small parties consisting of phenoty pically deviant cream-white, long-haired, fluffy-coated males of the Guianan Weeper Capuchin Ce. ( olivaceus ) castaneus. Such all-male parties seemed to range randomly while travelling at high speed through the vast landscape of pristine matrix lowland rain forest in the middle of which his study area was situated. It is located at more than one-hundred km north of Kaiser Mountains, a hilly country of which the foot- hills seem to form the Guianan Weeper Capuchin’s core distribution. This region that provides this monkey with its preferred habitat - ‘mountain savanna forest’- was found to sustain a very large population of this elsewhere in the Guianas ex- tremely rare taxon Ce. ( olivaceus ) castcmeus. Mountain savanna forest is typified by an under- story that is dominated by the majestic ‘bergi- maripa’ palm Attalea speciosa Mart. (Arecales A recaceae). A bove 400 m altitude, this palm tree is locally so abundant that one gets the impression to walk through a monocultural plantation of the African oil-palm Elaeis guineensis Jacq . (Arecales Arecaceae). The large fruits of Attalea speciosa constitute the Guianan Weeper Capuchin’s principal daily food throughout most of the year. Mountain savanna forest above 400 m altitude, therefore, may function as a ‘keystone habitat’ to the Guianan Weeper Capuchin, hence the high population density. Two decades later, while conducting biod- iversity surveys in Pico da Neblina National Park situated in the extreme northwestern corner of the Brazilian Amazon, the authors spotted a population of near-albinotic Weeper Capuchins that were characterized by a very dense, fluffy, overall long- haired, cream-white bleached fur. Their coat fea- tures looked very similar to that of the all-male parties that were seen sporadically passing through the Voltzberg study area. The Pico da Neblina population of weeper capuchins was spotted in a low type of cloud forest scrub that grows at high altitudes of 2,000 to 2,500 m. To the astonishment of the researchers, the capuchins were seen spend- ing part of the daytime on the ground in the middle of open tepui (sandstone table-mountain) ‘rock savanna’. They were seen foraging for inverteb- rates, mostly snails, other organisms endemic to tepui mountain tops, in addition to vegetable matter (e.g., roots, tubers and pseudobulbs of all sorts of terrestrial bromeliads and orchids). In retrospect, our sighting may be explained for as follows. In the past, a founder-colony of near-albinotic Guianan Weeper Capuchins, driven by the ‘trend to allo- patry’ out of the center of dispersion of archetypic Ce. ( olivaceus ) castaneus, may have traversed the upper Rio Branco and then reached the Pico da Neblina area. The latter is situated somewhat south of the Rio Cassiquiare, the channel that runs through the watershed connecting the basin of the Rio Negro with that of the Rio Orinoco. The fully bleached euchromic, long-haired, soft-coated weeper capuchins that were seen foraging in tepui cloud forest and open rock-savanna at 500-1,000 m below the 3, 004 m Pico da Neblina summitmuch resembled the near-albinotic, fluffy-coated Ce. ( olivaceus ) castaneus from Kaiser Mountains, Central Suriname. If the Pico da Neblina population turns out to represent a new taxon or one in-the- making, the ‘Neblina Weeper Capuchin’ would occupy a dead-end distribution in the southwestern- most corner of the Ce. olivaceus Clade’s range, the farthest away from the supposed center of the Clade’s dispersion (the Guianas or Venezuelan Coastal Range). The upper Rio Negro forms the di- vision between the distributions of the Guianan Weeper Capuchin Ce. olivaceus Clade and the Humboldt’s White-fronted Capuchin Ce. albiffflUS Clade (Fig. 39). This example from the field is in line with our theory of allopatric speciation in male- defended territorial primates such as CebuS. The ‘Neblina Weeper Capuchin’ may have radiated away from the Ce. olivaceus Clade’s center of dispersion in the Guianas following a pathway of metachromic bleaching driven by the trend to allo- patry in phenotypically deviant euchromic, long and flu ffy -h aired males. Interestingly, the mechanism of allopatric speciation and radiation of a mono- phyletic clade of monkeys like that of Humboldt’s Weeper Capuchins at first sight seems non-ad- aptive, at least in strict Darwinian sense, for it is solely based on discriminatory behavior performed exclusively by high-ranking males. The genes for warm, long and flu ffy -h aired coats are simply retained in the genes of these capuchin ‘founder- colonies’. Such a feature would therefore not a priori be the result of adaptive processes of natural selection. Its warm coat only secondarily happened to have survival value. It only turned adaptive when these gracile capuchins had to adapt in a short 178 Marc G.M. van Roosmalen & Tomas van Roosmalen period of time to a new habitat or feeding niche that would not have suited the species they derived from. Following this rational, one may speculate about a similar metachromic pathway that our hom- inid ancestors about 6 MYA must have followed when exchanging the canopy of tropical rain forest for a landscape of arid, open savanna scrub. Or a similar path way ofmetachromic bleaching towards albinotic (from a black to yellow or white skin color) and/or depilation of the body that different hominids followed between 100,000 and 50,000 years ago, when the trend to allopatry (male dis- criminatory behavior) forced them to leave the center of hominid dispersion and the cradle of hominid evolution - C and N Africa - to make a harsh living of nomadic big-game hunting/gath- ering in (for hominids) clime - and habitat - wise new, marginal, unsuitable, or inhospitable land- scapes of Central Europe, the Middle-East and SEA sia. Capuchin Monkeys of the genera CebllS and SapajuS formed distinct monophyletic Clades that diverged during the Late Miocene to Early Plio- cene, about 6.2 MYA. During the Plio-Pleistocene era the Clades diversified into two groups: Gracile or Untufted Capuchins genus CebllS, about 2.1 MYA in what is nowadays the western Amazon, and Robust or Tufted Capuchins genus ScipCljllS, begin- ning about 2.7 MYA in what are today SE Brazil, E Paraguay, and N Argentina. There is strong evid- ence from molecular genetic studies that Robust Capuchins (genus SapajuS) spent most of their evol- utionary history in the Atlantic Forest of SE Brazil, NE Argentina, and E Paraguay.And that the current wide-ranging sympatry of Robust and Gracile Capuchins across the larger part of the Amazon Basin is the result of a single, rapid, Late-Pleisto- cene invasion of Robust Capuchins from the At- lantic Forest, first into the ‘Cerrado’ and ‘Cerradao’ of C and NE Brazil, and only recently (about 0.4 MYA) from central South America north into the Amazon Basin and the Guianas (M it term eier et al., 2013). Though widespread throughout the Amazon Basin and the Guayanan Shield, the genetic differ- entiation of the Amazonian Robust Capuchins is limited. The fact that the phenotypic diversity of the Amazonian Robust Capuchins is not mirrored by a corresponding genetic diversity strongly supports our theory of allopatric prim ate speciation.A num- ber of the 16 taxa that are overall recognized in different taxonomic arrangements (e.g., Groves, 200 1 a; Silva Jr., 2001; Silva Jr., 2002) may well represent taxa ‘in-the-m aking ’ . Here, we follow Silva Jr. (200 1) in recognizing only two species: the Guianan Brown Capuchin Sap. apella with three subspecies distributed in the eastern Amazon and the Guianas, and the Large-headed Capuchin Sap. macrocephalus with four subspecies that are dis- tributed across the western Amazon. These taxa form two monophyletic Clades in which little genetic differentiation is shown. In contrast, the non- Amazonian species recognized by Silva Jr. are genetically distinct forming the monophyletic Sap. TligritliS Clade (Figs. 39, 40). Among the six species of the extra-A m azonian Sap. YligvitUS Clade we consider the Black-horned Capuchin Sap. nigvitUS the nearest to archetypic, less bleached species (Figs. 42, 43). Its southernmost populations repres- enting the darkest, overall m ost saturated eum elan in form may well be a distinct taxon named Sap. CUCullatUS by Spix in 1 823. The Black-horned Capuchin is the most S occurring of all robust capuchins. It is distributed in SE Brazil, S of the Rios Doce and Grande, extending S through the Atlantic Forest, and taxon Sap. CUCullatUS further south E of the Rio Parana into Rio Grande do Sul State and NE Argentina. The Black-horned Capuchin is a large-sized species with horn-like tufts on either side of the head at the temples. Its fur is overall very dark brown or grayish in nigritus, and black in Sap. CUCullatUS, often with slightly pheomelanin bleached, reddish or yellow-fawn colored underparts. A black to dark-grayish crown (with tufts in adults) contrasts much with the light colored face. The tail is black. From Sap. nigritUS derived the monotypic Crested Capuchin Sap. TO- bliStUS after a founder-colony of Sap. lligritUS tra- versed the Rio Doce to the north. It is distributed in SE Brazil from the Rio Jequitinhonha in Bahia State S to the Rios Doce and Suagui Grande in Es- pirito Santo State and E Minas Gerais State, E of the Serra do Espinhago. This taxon is very dark wood-brown or blackish above and on the limbs, with a faint dorsal stripe. The underparts are pheo- melanin bleached red or yellowish, whereas fore- arms, hands, lower legs, and feet are deep dark brown to black. Its face is dark grayish, with some white hairs on the forehead and temples. The crown tufts are tall and conical in shape. From a founder- colony of the northern Black-horned Capuchin Sap. On the origin of allopatric primate species 179 nigritUS that once traversed the Rio Jequitinhonha to the north, derived the Yellow -breasted Capuchin Sap. xanthoSterUOS. Yellow -breasted Capuchins tend to be much darker in overall color in the southwestern part (N Minas Gerais State), whereas they are pale in the northern part of this taxon’s dis- tribution. The monotypic taxon Sap . XaYlthoSteVYlOS is further distributed in CE Brazil, S and E of the Rio Sao Francisco, south to the Rio Jequitinhonha (in S Bahia State). It is generally pheomelanin bleached brindled reddish above with a sharply marked, golden-red underside. Tail and limbs re- mained saturated eumelanin black. Its crown does not contrast with the body, the cap is black, and the face and temples are fawn. It has small back- ward pointing tufts. From Yellow -breasted Sap. XanthoSterUOS derived the monotypic Blond Capuchin Sap. flavius, which was described by Schreber in 1774. Until it was collected in 2005, the Blond Capuchin was only known from an early illustration. Before colonial times, it must have been distributed in CoastalNE Brazil from S Rio Grande do Norte State through Paraiba State into NE Pernambuco. This taxon may extend its range to the left bank of the Rio Sao Francisco in Alagoas State. The Blond Capuchin is small, distinctive, and untufted. Its body and limbs are uniformly ad- vanced pheomelanin bleached g old en -y ello w , whereas its lower-body parts are slightly darker golden-yellow. Hands and feet are black, whereas the tail is uniformly golden-blond, but darker on the dorsal side than the rest of the body. It further has a rectangular snow-white cap on the front of the head, to just above the ears, and a furless, pendulous throat flap. Face and forehead are near-albinotic, cream to pinkish colored, the eyes are brown. SapajliS flaviliS occupies degraded CoastalAtlantic Forest and Montrichardia linifera (Arruda) Schott (Araceae) swamp in Pernambuco State, and ‘caatinga’ scrub in W Rio Grande do Norte State. Being advanced pheomelanin bleached to near- albinotic, taxon flavius occupies a dead-end distri- bution. It therefore fully concurs with our theory on the origin of allopatric speciation. The theory suggests that a founder-colony of progressively pheomelanin bleached Sap. XaYlthoSteVYlOS o nee was forced to make a living in the (for Robust Capuchins) marginal or unsuitable habitat of swamps and low xerophytic, spiny scrub of profusely branched bushy vegetation up to 8-10 m in height, mixed with prickly succulent cacti, and spiny, rigid-leaved bromeliads. Blond Capuchins are reported to use even sand dunes and mangroves. From the northern form of the Black-horned Capuchin derived to the W the Hooded Capuchin Sap. cay, and to the N the Bearded Capuchin Sap. UbidinOSUS (Fig. 42). The monotypic taxon cay is distributed in SE Bolivia, N Argentina, SW Brazil - W of the Rio Parana through Mato Grosso State into SW Goias and Mato Grosso do Sul - and Paraguay (E of the Rio Paraguay as far as the Rio Parana). The Hooded Capuchin Sap. cay is a small, short- limbed species without sexual dimorphism, typified mainly by its prominent dark dorsal stripe. SapajliS Cay is very variable in color, but generally rather pale. Its crown is pale to blackish -brow n , with two small hornlike tufts. Dorsal parts of the body (shoulders, front of the upper arms, saddle, rump, and thighs) are gray ish -b ro w n . Forearms, hands, wrists, lower legs, and feet are blackish. Eyes, nose, and mouth are surrounded by white hairs. It has a small white beard, and a dark line extends down from the ears to under the chin. From the Black-horned Capuchin derived to the north the monotypic Bearded Capuchin Sap. UbidinOSUS. This taxon is distributed in C and NE Brazil, W and N of the Rio Sao Francisco into Maranhao State, and in the W of Piaui State, and E to C Rio Grande do Norte, NW Paraiba, W Pernambuco, and W Alagoas; to the W it extends to the Rio Araguaia, and its southern limit is the north bank of the Rio Grande in Minas Gerais. To the west, the Bearded Capuchin taxon Sap. UbidinOSUS is replaced by Sap. apella, to the east by Sap. flavius, and to the south of the Rio Sao Francisco by Sap. Xanthostevnos . SapajliS nigritUS occurs just south of the Rio Grande. Some hybridization between Sap. Ubidi- UOSUS and Sap. nigritUS is reported to occur in the western part of Minas Gerais. The Bearded Capuchin Sap. UbidinOSUS is comparatively small and does not show sexual dimorphism. It differs from all other Robust Capuchins by the rusty-red hair on the back of the neck, the dark-brown preau- ricular stripe running down the side of the face in front of the ears, and the orange-yellow throat and dorsal parts of the body, flanks, outer part of arms, and proximal two-thirds of the tail. Forearms are dark, and the lower back and outer surface of thighs are gray ish -brow n , mixed with some reddish hairs. The crown is black, with rounded, sometimes bushy, black tufts. 180 Marc G.M. van Roosmalen & Tomas van Roosmalen Here, we recognize only two Amazonian Robust or Tufted Capuchins (genus Sapajus)-. the mono- typic Guianan Brown Capuchin Scip. Cipellci that is distributed in the eastern Amazon and in the Guianas, and the Large-headed Capuchin Scip. ITiaC- rocephalus with a number of forms/morphs/sub- species that are distributed throughout the western Amazon as far north as the Magdalena Valley in N Colombia (Fig. 42). Taxon Scip. Cipellci is found in the rain forests of the Amazon Basin ofBrazilN of the lower Rios Negro and Amazonas, E of the Rio Branco, extending N to the southeastern part of the Orinoco Delta in Venezuela and the Guianas. Its distributional limits in the S, SE, and E are defined by the extent of the Amazon rain forest, in the S and E of Maranhao State marking the transition zone to xeric deciduous forest and ‘caatinga’ scrub. In the West, its distribution is limited by the interfluve of the Rios Negro and Solimoes and the Rio Madeira Basin. The Guianan Brown Capuchin species Sap. apella is relatively large and heavily built, with a broad head, flat face, and short limbs. Its coat is long and coarse, with all five extremities darker colored than the rest of the body. It is generally gray-fawn to dark brown above, with a yellowish or red underside. The lower limbs and tail are black, and there is a variably distinct dorsal stripe. The face and temples are light gray-brown. The crown tuft is black and forms short tufts above the ears (the characteristic ‘horns’). The crown cap extends down the cheeks forming ‘sideburns’ that often meet below the chin. There is no sexual dimorph- ism, but males are slightly heavier and often overall much darker colored. The M argarita Island Capuchin taxon Sap. apella margaritae that is endemic to Isla de M argarita off the Caribbean coast of Venezuela fatutUus AMAZONIAN ROBUST CAPUCHINS SAPAJIS pallidum 1 union ns Sapajus apella Guianan Broun Capuchin ) 0 Sapajm apella margaritae Sapajus macracephalut Large- beaded Capuchin Colombian morph fatueilm o ■, Peruvian morph maranonh . ‘ Bolivian morph pallid us Brazilian morph juruanus Figure 44. Phylogeography, allopatric speciation. and metachromic bleaching in all Amazonian Robust Capuchins disputedly divided up in Sapajus Cipellci and S. maCWCephaluS, the latter with different taxa ’ in -the-m aking ’ . On the origin of allopatric primate species 181 distinguishes itself from the nominate Guianan Brown Capuchin by longer dark sideburns in front of the ears, and progressively bleached, pale-yellow or straw colored, near-albinotic upper arms and shoulders. The thighs and rump are pale yellow- brown, and flanks, lower back, and upper chest are pale brown, becoming paler from the upper back to the neck. The face is grayish, tinged pink on the cheeks and chin. The black cap extends in a “V” to between the eyes, with small round tufts above the eyes. The monotypic Large-headed Capuchin taxon Sap. macro cephalus is distributed in the western Amazon Basin, but its taxonomy and distributional limits are poorly defined. According to Silva Jr. (2001 ) this species includes the forms/morphs/sub- species Sap. fatuellus from the upper Magdalena Valley, Colombia, Sap. maraYlOYlis from Rio Ham- burgo, Peru, Sap. pallidus from the Rio B eni, C + N Bolivia, and Sap. jliruanus from the Rio Jurua, Brazil. Preliminary genetic studies in 2012 failed to indicate that Sap. apella and Sap. macrocephalus were distinct taxa. Large-headed Capuchins are distributed across the upper Amazon Basin in E Colombia, north as far as the Rio Arauca on the border with Venezuela, E Ecuador, E Peru, W Brazil, and C and N Bolivia (S at least as far as the upper Rio Beni). Their overall coat color is gray- brown or ochreous to dark brown above, with a dark dorsal stripe, and yellow-fawn or red-gold below. Sides of the neck are lighter, upper arms are pale yellowish, and legs are black with yellow-fawn or red-gold below. Adults have high, pointed crown tufts that resemble horns, which become reduced with age. There is often a gray-white stripe running from eye to ear. Four forms of the Large-headed RED NECKED GROUP uaucyntaae 0 Aotus miconax Aorus nigriceps Aotus a zame fa.) infulaim fa.) botivieniis fa.) azarae GREY-NECKED GROUP Aotus zonalis Aotus temunnus ( Aorus jorgeheruandezi # Aorus griseimembra Aotus brumbacki (ft Aotus vociprans Aotus tririrgatus Figure 45. Phy logeography, allopatric speciation, radiation, and metachromic diversification in all hitherto recognized taxa ofNight Monkeys, genus AotUS. 182 Marc G.M. van Roosmalen & Tomas van Roosmalen Capuchin have been distinguished (Fig. 44). The Colombian form Sap. fatuelluS is bright brown above and red below, having a prominent dorsal stripe. Its face is almost naked and dark-purplish to flesh-colored. The Peruvian form Sap. mavaYlOYlis is uniformly dark chestnut-brown above, becoming more reddish towards the flanks, and deep yellow- brown below. Its legs, tail, and (sometimes) fore- arms are black. Its cap is distinctly black, whereas temples and sides of the crown are often white. It has a crescent-shaped whitish patch above each eye. There are no crown tufts or they are minimal. The Brazilian form Sap. jUTUanilS is reddish-brown above with a very distinct blackish dorsal stripe. The throat and upper chest are blackish or pale reddish-buff, and limbs and tail are dark brown or black. The Bolivian form Sap. palliduS fro m south of the Rio M adre de Dios has also been referred to as a subspecies of Sap. libidinOSUS, but such tax- onomy would be conflicting with our theory on al- lopatric speciation, for Sap. libidinOSUS from CE Brazil is overall more advanced pheomelanin bleached in comparison with Sap. palliduS. Both the Colombian morph/taxon Sap. fatuelluS of the Large-headed Capuchin Sap. maCWCephalus and the insular Margarita Island Brown Capuchin Sap. apella rnargaritae are in their overall advanced pheomelanin bleached coat coloration clearly fol- lowing the metachromic pathway to albinotic, and therefore fully concur with our theory of allopatric speciation (Fig. 44). NightMonkeys orDouroucoulis genus AotUS rep- resent a very old lineage that is generally placed in a family of its own - Aotidae. The molecular genetic evidence classifies them as a subfamily of the Ce- bidae. There is also morphological evidence to place AotUS in the Pitheciidae. There are generally two Groups distinguished: the “Gray-necked Group” (characterized by grayish to brownish agouti sides of the neck and body), which occurs north of the Amazon River, and the “Red-necked Group” (characterized by partly or entirely orange or yellowish sides of the neck and chest, much contrasting with the grayish to brownish-agouti colored sides of the body), which occurs south of the Amazon River (Mittermeier et al., 2013). Re- cently, up to eleven species have been recognized, of which at least seven in the Gray-necked Group: the Lem urine Night Monkey Ao. leiflUritlUS, the Pan am anian Night Monkey Ao. ZOTialis, B rum back’s Night Monkey Ao. bvuvnbacki, the Gray-legged Night monkey Ao. griseiffieiTlbra, Spix’s Night M on key Ao. VOciferans, Humboldt’s Night Monkey Ao. trivirgatUS, and Hernandez-Camacho’s Night Monkey Ao. jorgehemandezi (Figs. 45, 46). In the Red-necked Group are recognized four species: the Andean Night Monkey Ao. JfliconaX, Ma’s Night Monkey Ao. YiancyiTiaae, the Black-headed Night M onkey Ao. VligricepS, and A zara’s N ight M onkey A. azarae (Figs. 45-47). Sexual dimorphism in night monkeys is absent. They are also not sexually dichromatic in coloration and facial markings. The coat is in metachromic sense primitive, archetypic saturated eumelanin, grayish to grayish-tan with a pheomelanin bleached, lighter tan or yellowish underside. In Red-necked species, ventral surfaces of neck, chest, abdomen, and inner sides of arms and legs are orangish or russet colored. The faces have white patches over eyes, topped by black stripes, and a triangular black patch running from the center of the forehead down between the eyes. Black stripes are also extending from the lateral side of each eye to the forehead, varying in width and darkness, and may or may not converge posteriorly with the central stripe. Tails are generally agouti- brown, distally black-tipped. Night Monkeys most likely descended from a diurnal haplorrhine. They only are secondarily nocturnal and have retained their color vision . Within the Gray-necked Clade Ao. letnurinus is the nearest to archetypic, saturated eumelanin, less bleached taxon. It is a montane species of the Colombian Andes range, at elevations above 1,000- 1,500 m, in the upper Rio Cauca Valley and on the slopes of the Cordillera Oriental (but not in the Magdalena Valley that is occupied by the Gray- legged Night Monkey Ao. griseimetnbra), extend- ing its range S into Ecuador through the humid sub- tropical forests of the Cordillera Oriental. The Lem urine Night Monkey is rather shaggy and long- haired, with the upperparts of the body often eu- melanin grayish to buffy-agouti, with a poorly defined brownish medial dorsal band. The under- side of the body is pheomelanin bleached yellowish to pale orange. Inner and outer sides of limbs are entirely grayish-agouti, or the inner sides have a yellowish to pale orange tone extending from the chest and belly to the mid-arm or mid-leg. Hands and feet are dark. Temporal stripes may be separ- ated or united behind the head. From ancestral Ao. On the origin of allopatric primate species 183 lemurinus derived the Gray-legged Night Monkey taxon Ao. griseimembra. It is distributed in N Colombia and NW Venezuela. It occurs in the Rio Magdalena Valley and northern lowland forests of Colombia (including the Sierra Nevada de Santa Marta and the Rios Sinu and San Jorge basins), extending into Venezuela in the vicinity of Lake Maracaibo. It is grayish to brownish-agouti on the side of the neck. Upperparts are grayish to buffy; chest, belly, and inner surfaces of the legs are brown- ish or yellowish to pale orange. Pelage is relatively short. Hands and feet are light-brown. From taxon Ao. griseimembra derived to the NW the mono ty pic Panamanian Night Monkey zonalis. This taxon is distributed in NW Colombia in the Pacific low- lands, S towards the Ecuadorian border, and W into most of Panama; it is absent from SW Panama (Chiriqui). Its overall coat color is brownish in the Canal Zone and Colombia, but it grades into paler and grayer tones along the upper Rio Tuira, E Panama. From Ao. ZOYialis derived Hernandez- Camacho’s Night Monkey Ao. jorgehernandezi. This monotypic taxon is believed to occur in the (sub)-m ontane tropical forests on the western slopes and foothills of the W Colombian Andes (in Quindio and Riseralda). It is advanced bleached to albinotic in the head and ventral parts. Its face has two discrete supraocular white patches separ- ated by a broad black frontal stripe. Moreover, subocular white bands of fur are separated by a thin black malar stripe on each side of the head. Ventral parts of the arms from the wrists running up into the chest and belly are of a thick white fur (Fig. 39). From the Gray-necked Night Monkey Clade’s nearest to archetypic taxon Ao. lemurinus derived to the SE first Brumback’s Night Monkey Ao. brumbacki. This monotypic taxon is distributed in NC Colombia in the eastern part of Boyaca De- partment, E to the highlands of Meta (to at least 1,500 m above sea level). Its coat is dorsally grayish-buffy agouti colored with a dark brown mid-dorsal zone. Ventral parts extending to the elbows, knees, and lower throat are pale orange. Sides of the neck are entirely grayish or brownish agouti, like the flanks and outer sides of the arms. The head shows well-marked, thin, brownish-black temporal stripes. The white above the eyes is yel- lowish, and the white on the face extends to the chin. From Ao. brumbacki derived first to the S Spix’s Night Monkey Ao. VOciferans. This mono- typic taxon is widespread in the upper Amazon Basin, extending from NW Brazil (W of the Negro, Uaupes, and A m azon as-S olim 5 es Rivers) into SE Colombia (S of the Rio Tomo, Orinoco Basin), and S into the Ecuadorian Amazon and NE Peru (as far south as the north bank of the M aranon-Am azonas River). It occurs also S of the Rio Solimoes in a small area on the lower Rio Purus. Spix’s Night Monkey’s coat is brown-toned above, with an over- all white, slightly orange tinged underside, extend- ing to the wrists, ankles, and chin. Hands and feet are black. The proximal one-third to one-half of the ventral side of the tail is reddish or grayish- red, the rest is black. The crown stripes on the head are thick and brownish, with white fur above the eyes con- fined to two small patches grading into the agouti- colored crown. The temporal stripes are united behind, and the malar stripe can be well defined to absent. The face is white, except for the chin. From Ao. VOCiferailS derived to the N and E the mono- typic Humboldt’s Night Monkey taxon Ao. trivir- gatUS. It is widespread across N Brazil, N of the Rios Negro and Amazonas and W of the Rio Trombetas, N into SC Venezuela and E Colombia. Sides of the neck are grayish-agouti to mainly brownish-agouti colored. Upper parts of the body are grayish to buffy-agouti. The inner sides of the limbs, extending to the wrists and ankles, are sim- ilar in color to the orange-buffy of chest and belly. The face has triradiate brown stripes. It is rather grayish in comparison with the usual white of other Night Monkeys. Hands and feet are dark-brown. Taxon Ao. trivirgatUS can be distinguished from all other Night Monkeys by its parallel temporal stripes on the head and the lack of an interscapular whorl or crest (Figs. 45, 46). Within the Red-necked Clade of Night Mon- keys, the Black-headed Night Monkey Ao. Yligri- ceps is the nearest to archetypic, less pheomelanin bleached taxon (Fig. 47). This monotypic species is distributed in the Brazilian Amazon, S of the Rio A m azo n as-S o lim oes and W of the Rio Tapajos- Juruena, as far south as the right bank of the Rio Guapore and the left bank of the Rio Madre de Dios in N Bolivia. It occurs also in SE Peru, west to the Rio Huallaga, and north as far as the Rio Cush- abatay. Its coat is iron-gray above and brownish- agouti on the dorsum. The underside is orange colored with white tones, extending to the neck, throat, chin, and sides of the jaw and also to the 184 Marc G.M. van Roosmalen & Tomas van Roosmalen GRAY-NECKED NIGHT MONKEYS AOTUS Figure 46. Radiation and metachromic d iversification in the Gray- necked Night Monkey Group ( AotUS ), folio w in g eum elan in and pheomelanin pathways of metachromic bleaching, in particular in the head, proximal half of the tail, and ventral parts of the body. On the origin of allopatric primate species 185 Figure 47. Radiation and metachromic diversification in the Red-necked Night Monkey Group ( AotUS ), following eu - melanin and pheomelanin pathways of metachromic bleaching, in particular in the head, tail, and ventral parts of the body. 186 Marc G.M. van Roosmalen & Tomas van Roosmalen inner surfaces of the wrists and ankles. The cap is black, the face stripes are broad, and it has distinct areas of white on the face. From the Black-headed Night Monkey Ao. nigri- ceps derived to the W Ma’s Night Monkey Ao. TICincy lTiaae . This monotypic taxon ranges in W Brazil (S of the Rio Solimoes from the Rio Javarr as far east as the Rio Jandiatuba) and NE Peru (from the Rio Javari W to the Rio Fluallaga). This taxon is also found in an enclave between the lower Rios Tigre and Pastaza. The upper parts of its coat are grayish-agouti, with a dark mid-dorsal zone and a pale orange underside, extending up the sides of the neck and inner limbs. The proximal part of the tail is orange, with a blackish stripe above; the under- side is blackish. Its face is grayish-white, the crown stripes are narrow and dark brown colored, and the sides of the throat and jaw are colored like the body (Fig. 41). From Ao. nancymaae derived to the W the Andean Night Monkey taxon Ao. Uliconax. This monotypic night monkey is endemic to Peru. It is confined to a small area S of the Rio Maranon and W of the Rio Huallaga. It inhabits the primary and secondary humid, lower-montane cloud forests in the Andes at elevations of 800-2,800 m. Upper sides of its coat are light gray with a brownish tint, often quite infused with red-brown. Its underside is pale orange, extending forward as far as the chin and on the inner sides of the limbs. Outer surface of the body is overall brownish to buffy-agouti. The tail is bushy, its upper side is blackish, its lower side reddish-orange. Head parts and throat are advanced bleached to near-albinotic . From the nearest to archetypic Red-necked Black- headed N ight M onkey Ao. nigriceps derived in op- posite direction (to the S and E) Azara’s Night Monkey species Ao. azarae. Three subspecies of Ao. azarae are recognized: the nominate taxon Ao. azarae, distributed in SC Brazil, S Bolivia, Paraguay, and N Argentina; taxon Ao. boliviensis, distributed in SE Peru and Bolivia east of the Andes; taxon Ao. infulatUS, distributed in Brazil, S of the Rio Amazonas (but with a small enclave in the SE tip of Amapa State), including M arajo and Caviana Islands, extending east into Maranhao State as far as the Rio Parnalba, S along the west bank of the Rio Tocantins to the Pantanal of Mato Grosso. Taxon Ao. azarae inf Hiatus’ s western limit is the Rio Tapajos-Juruena.Azara’s Night M onkey Ao. azarae is highly variable. It generally has an in- terscapular whorl. Taxon Ao. azarae has a long, thick, and shaggy fur that is grayish to pale buffy- agouti above and pale w hitish -oran ge below. Facial stripes are narrow. The basal hairs of the distal l A of the tail are orange. Taxon Ao. boliviensis has a relatively short fur, with an olive tone above and contrastingly grayer on the limbs. The facial stripes are very narrow except where the middle one ex- pands on the crown; the black temporal stripe in this taxon is poorly defined, the black malar stripe is faint or absent, and there is usually a whitish band between the eyes and temporal stripe. There is a conspicuous whorl between the shoulder blades. The third taxon Ao. infulatUS, the “Feline Night Monkey”, is very similar to subspecies Ao. bolivi- ensis, but the white on the face is more prominent. There is no whitish band between the eyes and the temporal stripe as there is in Ao. boliviensis. The temporal stripes are black, well defined, and con- tinuous with the malar stripe. The tail is reddish throughout its length except for the black tip. The orange color of the underparts extends to or above the ventral one-half of the sides of the neck. The color of the throat varies from orange, with the anterior one-half grayish-agouti to entirely orange colored (Fig. 47). Our theory suggests that the trend to allopatry in Neotropical primates resulted from a specific kind of social selection. That the discrimination of somewhat deviant mutant young males by high- ranking males, which push them toward the peri- phery of the parental group’s range, has been the true driver behind metachromic bleaching on the evolutionary path along which a certain race, species, phylogenetic clade, or genus has extended its geographic range in the past. As any founder- colony or population at the limit of a taxon’s current range will represent a narrow gene pool, through in- breeding certain phenotypic characters (e.g., local depilation of the skin, change of coloration of the skin, pelage or parts of it) will initially be reinforced and advance more rapidly within the population. Through the process of metachromism (changing hair and skin color) with the trend to allopatry as the behavioral driving force, speciation, radiation, and phylogeography can be retraced and well explained for in all extant Neotropical primates. According to the principle of metachromic bleach- ing, extant primate taxa at the base of a phylogen- etic tree or clade are in general agouti or saturated On the origin of allopatric primate species 187 eumelanin colored. They are the least colorful, black(ish) or dark brown toned, and therefore con- sidered to be nearest to the ancestral, archetypic, primitive or original form. Geographic variation and diversification in color patterns of the coat among Neotropical mon- keys demonstrates with unusual clarity the unilat- eral direction and irreversibility of processes that lead to progressively metachromic bleached and ultimately (near)-albinotic allopatric forms, irre- spective of environmental factors (Figs. 1-47). The essentially behavioral and genetic driving forces behind metachromic processes, though, have never been studied. They are generally considered enig- matic. The reason may be that they seem to disobey commonly accepted Darwinian rules of evolution. Different from birds, in territorial (Neotropical) monkeys metachromic changes in coat color toward bleaching or albinotic and/or all sorts of local hair growth or loss of hair (depilation) do not seem to play an essential role in sexual display and mate selection. Consequently, they may seem to be non- adaptive. In the wild only rarely one is able to witness how exactly processes of metachromic bleaching do work out. For instance: when some- what bleached or depilated deviant young males are being pushed from the center into the periphery of a ranging or foraging group. Or: when ‘outcast’ males do join in all-male parties. Or: when such parties set out to look beyond the horizon, for mere survival willing to overtake any habitat delimitation or geographic barrier found on their ‘path to allo- patry’. These crucial data will only come available when fieldw orkers, like we did, do live for pro- longed periods of time among undisturbed primate populations in pristine tropical forest environment. As very few prim atologists have done so, at least in the Neotropics, and sample sizes are consequently too small to be published and divulged, it is im- possible for us to add more references then our own on the matter. Even though, living over more than a decade in permanent intimate contact with pristine nature, both in the Brazilian Amazon and in the over- all even better preserved Guayanan Shield, led us to believe that high-ranking males pushing slightly bleached and/or depilated young males to the peri- phery of a group’s range, or sometimes beyond its boundaries, could plausibly be the true and prin- cipal motor or driver behind allopatric speciation and radiation of taxa in nearly all Neotropical primate genera - Pygmy Marmosets ( Cebuella . ), Tamarins ( SdguiriUS ), Amazonian Marmosets ( Mico ), True Marmosets ( Cdllithfix ), Lion Tamar- ins ( Leontopithecus ), Sakis ( Pithecia ), Bearded Sakis ( Chiropotes ), Uakaris ( CacajaO ), Titi Mon- keys (CaUicebuS ) , Night Monkeys ( AotliS ), Squirrel Monkeys ( Sciiffliri ) , G racile/U ntu fted Capuchin Monkeys ( CebliS ), Robust/Tufted Capuchin Mon- keys ( SapajUS ), Howling Monkeys ( Alouattd ), Woolly Monkeys {LagOthviX ) , Spider Monkeys ( AteleS ), and Woolly Spider M onkeys (BrachyteleS) . Interestingly, but concurring with our theory (for those monkeys that do not defend a common ter- ritory), metachromic bleaching did not take place in peaceably living monkeys like the archetypic agouti and saturated eumelanin colored Black- crowned Dwarf Marmosets Cdllibellci huiflilis, a newly identified monotypic genus of diminutive callitrichid monkeys (Figs. 2, 3). Nor did it take place in saturated eumelanin all-black Goeldi’s Monkeys {CdllimicO goeldii) - the only other mono- typic primate genus in the Neotropics that does not behave territorial in any sense and therefore does not defend a common living space against the neigh- bors of its own kind (Fig. 4). Their external features showing archetypic agouti and saturated eumelanin coat coloration without any sign of metachromic bleaching are in full accordance with their genetics that put them at the base of their respective phylo- genetic trees. It further corroborates our theory on the origin of allopatric speciation in primates and the principle of metachromic bleaching, for Dwarf Marmosets and Goeldi’s Monkeys are equally so- ciable, peaceable little m onkeys that do not demon- strate any rate of territorial defense. The primitive agouti and saturated eumelanin (blackish-brown) Black-crowned D w arf M arm oset stands at the base of the phylogenetic tree of all Amazonian marmo- sets (Van Roosmalen & Van Roosmalen, 2003). It represents the nearest to ancestral, archetypic mar- moset from which all extant, advanced and highly territorial Amazonian marmosets (genus MicO ) and pygmy marmosets (genus Cebuella) have derived in the Late Pleistocene. Our theory is firmly rooted in over 30-year field- work on primates, both in the Guianas and in the entire lowland Amazon Basin. Lrom the very be- ginning we have given special attention to issues like socio-ecology, ecological feeding niches, ter- ritorial behavior, distributions, and phylogeography. 188 Marc G.M. van Roosmalen & Tomas van Roosmalen Simultaneously, we have kept, raised, bred, rehab- ilitated, and reintroduced back into the wild entire families or social groupings of a multitude of mon- key taxa representing about all hitherto known Neo- tropical primate genera. Many unique, extremely rare or sometimes once- in -a-lifetim e observations that we gathered in pristine tropical rainforest en- vironment as well as in captivity (the bulk of it never published inherent to ‘insignificant’ sample sizes) now do add up to the validity of our theory. It basically helps us to better understand the com- plex distribution patterns, phylogeography, diversi- fication, speciation, and radiation in Neotropical primates. Most likely, the theory applies to all the world’s primates (including man), as long as the taxa exhibit social groupings that defend a common living space, home range, or territory. The fact that only two Neotropical primate genera ( CcilluflicO and Callibella ) are monotypic strongly supports our theory, as it does not apply to peaceable, non- territorial social primates. By boat, canoe, and on foot we have surveyed entire basins of a number of major tributaries of the mighty Amazon River to study primate diversity and distributions across the entire Amazon Basin, including also large parts of the Brazilian and Guyanan Shields. We have tested and empirically come to fully validate Alfred Rus- sel Wallace’s river-barrier hypothesis that he first laid down in his 1 852 account On the Monkeys of the Amazon, and later in his 1876 paper “The Geo- graphical Distributions of Animals”. Herein, Wal- lace points at the larger rivers that he sailed as the principal evolutionary cause of the Amazon’s rich extant primate diversity and complex biogeography, since many rivers effectively block off gene flow between populations along opposite riverbanks (genetic isolation). As the Amazon still represents a largely pristine and vast natural realm that is (not yet) drastically and irreversibly modified by human interference, no better place to study and retrace evolutionary processes that may have acted upon primates and other mammals since the Pliocene era, no matter on which continent. Moreover, most rivers that in the course of millions of years have played a significant role in the demography of Amazonian primates - the majority of which cannot swim or fly- remain acting as such. Therefore, distributions of primate taxa in the Amazon, if correctly studied, documented, and taxonom ically treated, do follow a more transparent and rational overall pattern in comparison with those of the Old World. In SE Asia, instead of rivers, the ocean played an equally important role in the island bio- geography of mammals. And in Africa (including Madagascar), the landscape with its complex and diffuse mosaic of vegetation types and habitats seems to have played a more determinant role than rivers in primate distributions. Moreover, massive human disturbance has long irreversibly changed the landscape of the Old World. This may have obscured to some extent the principal factors that influenced and determined distributions and phylo- geography in catarrhine primates, most importantly the horn inins. CONCLUSIONS Here we discuss the above proposed doctrine on the origin of allopatric primate species and the prin- ciple of metachromic bleaching among Neotropical primates as a conclusive socio-ecological answer to the question: why primates are such a highly diver- sified, species-rich, and colorful order in the Class Mammalia. The Order Primates contains a world total of 73 genera, 414 IU C N -recognized species, and 612+ known taxa of which roughly one third are found in the Neotropics (Mittermeier et al., 2013) (see also Table 1). Globally, only the rodents (Order Rodentia) outnumber the Order Primates. However, compared to primates, rodents are by far not that diversified. They are mostly opportunists, not very sociable, and not particularly colorful. While studying color variation in c a llitric h id mon- keys, Hershkovitz (1968; 1977) pro posed the “Theory of M etachrom ism .”. He attributed evolu- tionary change in mammalian tegumentary colors to social, sexual, and predatory selection, as it seems to be the case in birds. He argued that the highly ‘visually’ adapted primates may be predis- posed to select mates based on coat color and hair adornments. However, primates generally do not sexually display their skin and coat colors, or hair dresses, except for a few genera in the Old World (e.g., Theropithecus, Mandrillus). instead, some display their genitals, like both sexes of Amazonian Marmosets (genus Mico ) do. Or, both sexes of Bearded Sakis ( ChiwpOteS ), female Spider Mon- keys (Ateles) or male Woolly Spider Monkeys ( Brachyteles ) do. In that case, their genitals are On the origin of allopatric primate species 189 mostly hypertrophied (e.g., MicO, CHiwpOteS, Bva- chyteles. Pan). Hershkovitz’s key hypothesis of m etachrom ism , which is tested in tamarins (genus SaguinuS) and confirmed for many of its predictions by Jacobs et al. (1995), concerns the orderly, irre- versible loss of pigment within chrom ogenetic fields. Its key concept is that genetic drift together with social selection could fix phenotypes departing from primitive agouti or saturated eumelanin (blackish-brown) fields by various degrees of so-called “metachromic bleaching”. Thus, an al- binotic (nearly white) coat would represent the end point of geographic variation in a series of near-al- lopatric forms (color morphs) deriving ultimately from an agouti-colored or saturated eumelanin pig- mented ancestral form. Using m etachrom ism , we have demonstrated that most Amazonian monkey genera are monophyletic and composed of two or more major phylogenetic Groups or Clades. We found only two genera (i.e., Callibella and Cal- limico) to be monotypic. Contrary to Hershkovitz, who followed the Darwinian fallacy of adaptive evolution by linking evolutionary change in mam- malian tegumentary colors (‘bleaching’) to social, sexual, and predatory selection, we suggest to at- tribute metachromic diversification in extant social and territorial prim ates exclusively and uniquely to “male social selection”. We propose the “trend to allopatry in somewhat metachromic bleached and/or depilated varieties” to be the principal mechanism and driver behind speciation, radiation, and phylogeography in group-living Neotropical monkeys that defend the group’s living space. It arguably applies also to any group-living territorial primate worldwide, including our own species and its ancestors (be it hominids or hominins). For all nineteen genera of Neotropical primates we have presented distribution maps of all known extant taxa and indicated the geographic barriers (rivers, lakes, mountain ranges, seasonally inundated flood- plain forests, open scrub areas, etc.) delineating each taxon’s distribution. We have also elaborated the phylogeography and radiation within each monophyletic cladistic Group or Clade and related them to the irreversible patterns of metachromic bleaching. Through the process of metachromism (changing hair and skin colors) with the trend to al- lopatry as the behavioral driving force, speciation, radiation, and phylogeography can be well retraced and explained for in all extant Neotropical primates. According to the principle of metachromism, prim- ate taxa at the base of a phylogenetic tree or clade are in general agouti or saturated eumelanin (black or blackish-brown) colored - that is the least color- ful. Within that Clade they are considered the nearest to ancestral, archetypic, primitive, or ori- ginal taxon. Based on metachromic skin and fur characters, without a single exception, we were able to retrace phylogeographic pathways of speciation and radiation that were plausibly followed in the evolutionary history of each monophyletic Clade. In all cases we could confirm the trend to allopatry following irreversible eumelanin and pheomelanin pathways of metachromic bleaching. The farther a taxon radiated away from the origin or center of the Clade’s dispersion, the more progressively eu- chromic or bleached and eventually albinotic its coat/pelage, or part of it, will become. The great majority of primates are sociable, group-living animals. Group sizes vary from nuc- lear families (4-7 individuals) to troops of mixed age and sex classes containing 15 to over 200 indi- viduals. The far majority of the world’s primate so- cieties are socially structured in a hierarchic way and based on male dominance and ranking. Male defense of the group and its living space within a population benefits from male social selection. Even in m atriarchally organized social groups, such as those of spider monkeys ( Ateles ) and pygmy chimpanzees or bonobos {Pan), males associate in all-male parties to jointly patrol and defend the group’s territory or living space. In social conflicts among males over ranking, inferior males as well as mutant males that show somewhat different, deviant phenotypic characters (such as a slightly bleached pelage here or there or depilated skin in certain body parts) will be pushed into the periphery of the group during ranging and foraging. We have seen this happening, both in the wild and in semi- free ranging conditions, in particular in social groups and societies of monkeys like LagOthrix, Ateles, Cebus, Sapajus, Saimiri, Cacajao, or Chiro- potes. Depending on the species, such young males also happen to be expelled from the parental group. We have witnessed this in wild and semi-free ranging populations of Alouatta, CalUcebuS, MicO, Cebus, Sapajus, and Pithecia. Either way, the chances of outcast males to survive and pass on their mutant genes are utterly slim. If this would happen in other mammals - being comparatively 190 Marc G.M. van Roosmalen & Tomas van Roosmalen Alouatta Lacepede, 1799 Alouatta arctoidea Cabrera, 1940 Alouatta belzeblll (Linnaeus, 1766) Alouatta caraya (Humboldt, 1 8 1 2) Alouatta discolor ( s p ix , 1 8 2 3) Alouatta guariba guariba (Humboldt, 1 8 1 2 ) Alouatta guariba clamitans Cabrera, 1940 Alouatta macconnelli Elliot, 1 9 1 o Alouatta nigerrima Lon n berg, 1 9 4 1 Alouatta palliata palliata (Gray, 1 8 4 8) Alouatta palliata aequatorialis Festa, 1903 Alouatta palliata coibensis Thomas, 1902 Alouatta palliata mexicana m erriam , 19 02 Alouatta palliata trabeata Lawrence, 1933 Alouatta pigra Lawrence, 193 3 Alouatta sara Elliot, 1 9 1 0 Alouatta seniculus seniculus (Linnaeus, 1766) Alouatta seniculus juara Elliot, 1 9 1 0 Alouatta seniculus puruensis Lonnberg, 1 9 4 1 Alouatta ululata e ilio t, 1 9 1 2 AotUS Illiger, 1811 Aotus azarae azarae (Humboldt, 1 8 1 2 ) Aotus azarae boliviensis Elliot, 1907 Aotus azarae injulatus ( k u h l, 1 8 2 o ) AotUS brumbacki Hershkovitz, 1983 Aotus griseimembra E liiot, 1912 Aotus jorgehernandezi Defier et b ueno, 2 00 7 Aotus lemurinus I. Geoffroy Saint-Hilaire, 1843 Aotus miconax Thomas, 1927 Aotus nancymaae Hershkovitz, 1983 Aotus lligriceps D o 11m a n , 19 09 Aotus trivirgatus (Humboldt, 1811) Aotus vociferous (Spix, 1 82 3) Aotus zonalis Goldman, 1914 Ateles E. Geoffroy Saint-Hilaire, 1806 Ateles belzebuth E . Geoffroy Saint-Hilaire, 1806 Ateles chamek (Humboldt, 1 8 1 2 ) Ateles fusciceps fusciceps Gray, 18 65 Ateles fusciceps rufiventris Sciater, 1872 Ateles geoffroy i geoffroyi k u h l , 1 8 2 o Ateles geoffroyi azuerensis (Bole, 1937) Ateles geoffroyi frontatus (Gray, 1842) Ateles geoffroyi grisescens Gray, 1865 Ateles geoffroyi ornatus (Gray, 1 8 7 o ) Ateles geoffroyi vellerosus Gray, 1865 Ateles geoffroyi yucatanensis Kellogg et Goldman, 1944 Ateles (hybridus) hybridus I. Geoffroy Saint-Hilaire, 1829 Ateles (hybridus) br urine us G ray, 1870 Ateles longimembris Allen, 1 9 1 4 Ateles marginatus E. Geoffroy Saint-Hilaire, 1809 Ateles paniscus (Linnaeus, 1758) Brachyteles Spix, 1823 Brachyteles arachnoides (E. Geoffroy Saint-Hilaire, 1806) Brachyteles hypoxanthus (K u hi, 1 8 2 o ) Cacajao Lesson, 1840 Cacajao (calvus) calvus ( I. Geoffroy Saint-Hilaire, 1847) Cacajao (calvus) novaesi Hershkovitz, 1987 Cacajao ( calvus) rubicundus (I. Geoffroy Saint-Hilaire et D eville, 1848) Cacajao ( calvus) ucayalii (Thomas, 1928) Cacajao ( melanocephalus ) melanocephalus (Humboldt, 1812) Cacajao (melanocephalus) ayresi Boubli, Silva, Hrbek, Pontual et Farias, 2008 Cacajao (melanocephalus) hosomi Boubli, Silva, Hrbek, Pontual et Farias, 2008 Cacajao ouakary ( s p ix , 1 8 2 3) Callibella van Roosm alen M .G .M . et van Roosm alen T., 2003 Callibella humilis (van Roosmalen M .G.M ., van Roosmalen T., Mittermeier et de Fonseca, 1998) Callicebus Thomas, 1903 Callicebus aureipalatii Wallace, Gomez, Felton A. et Felton A.M., 2006 Callicebus baptista Lonnberg, 1939 Callicebus barbarabrownae Hershkovitz, 1990 Callicebus bernhardi van Roosmalen M.G.M., van Roosm alen T. et Mittermeier, 2002 Callicebus brunneus (Wagner, 1842) Callicebus caligatus (Wagner, 1842) Callicebus caquetensis Defier, Bueno et Garcia, 2010 Callicebus drier ascens ( s p ix , 1 82 3) Callicebus coimbrai Kobayashi et Langguth, 1999 Callicebus cupreus ( s p ix , 1 8 2 3) Callicebus donacophilus (d ’ o rb ig ny, 1 8 3 6) Callicebus dubius Hershkovitz, 1988 Callicebus hoffmannsi Thomas ( 1 9 o 8 ) Callicebus lucifer Thomas, 1914 Callicebus lugens (Humboldt, 1812) Callicebus medemi Hershkovitz, 1963 Callicebus melanochir ( Wied-Neuwied, 1820) Callicebus modestus Lonnberg, 1939 Callicebus moloch (H offm annse gg, 1807) Callicebus nigrifrons (Spix, 1 823) Callicebus oenanthe Thomas, 1924 Callicebus olallae Lonnberg, 1939 Callicebus ornatus (Gray, 1866) Callicebus pallescens Thomas, 1907 Callicebus personatus (E . Geoffroy Saint-Hilaire, 1812) Callicebus purinus Thomas, 1927 Callicebus regulus Thomas, 1927 Table 1/1. References of scientific descriptions of all known Neotropical primates (present paper). On the origin of allopatric primate species 191 Callicebus stephennashi van Roosm aien m .g .m van Roosm ale n T. et M ittermeier, 2002 Callicebus torquatus ( H o ffm an ns egg, 1807 Callicebus vieivai Gualda-B arros, Nascimento et Amaral, 2012 Callimico Miranda Ribeiro, 1912 Callimico goeldii (Thomas, 1904) Callithrix Erxleben. 1777 Callithrix aurita (E . Geoffroy Saint-H ilaire, 1812) Callithrix flaviceps (Thomas, 1903) Call ithrix geoffroy i (Humboldt, 1 8 1 2 ) Callithrix jacchus (Linnaeus, 1 758) Callithrix kuhlii Coimbra-Filho, 1985 Callithrix penicillata ( E. Geoffroy Saint-H ilaire, 1812) Cebuella Gray, 1865 Cebuella ( pygmaea) pygmaea ( s p ix , 1 82 3 ) Cebuella (pygmaea) niveiventris Lonnberg, 1940 Cebus Erxleben, 1777 Cebus aequatorialis a lien , 1 9 1 4 Cebus albifrotlS (Humboldt, 1 8 1 2 ) Cebus brunneus a lien , 1 9 1 4 Cebus capucinus capucinus (Linnaeus, 1758) Cebus capucinus curtus Bangs, 1905 Cebus cesarae Hershkovitz, 1949 Cebus cuscinus Thomas, 1901 Cebus imitator Thomas, 1903 Cebus kaapori Q u e iro z , 19 9 2 Cebus leucocephalus Gray, 18 65 Cebus malitiosus Elliot, 1909 Cebus olivaceus olivaceus Schomburgk, 1848 Cebus olivaceus castaneusl. Geoffroy Saint-Hilaire, 1851 Cebus unicolor s p ix , 18 2 3 Cebus versicolor Pucheran, 1845 Cebus yuracus Hershkovitz, 1949 Chiropotes Lesson. 1840 Chiropotes albinasus (I. Geoffroy Saint-Hilaire et Deville, 1 848) Chiropotes chiropotes (Humboldt, 1812) Chiropotes sagulatus (Traill, 1 8 2 1 ) Ch iropotes sat anas ( H o f f m a n n s e g g , 1 8 0 7 ) Chiropotes Utahickae Hershkovitz, 1985 LdgOthrix E. Geoffroy Saint-Hilaire, 1812 Lagothrix (cana) cana ( E. Geoffroy Saint-Hilaire, 1812) Lagothrix ( cana ) tschudii Pucheran, 1857 Lagothrix ( lagotricha > lagotricha (Humboldt, 1 8 1 2 ) Lagothrix (lagotricha) lugens Elliot, 1907 Lagothrix poeppigii Schinz, 1844 Leontopithecus Lesson, 1840 Leontopithecus caissara Lorini et Persson, 1990 Leontopithecus chrysomelas (K u h l, 182 0) Leontopithecus chrysopygus (M ik a n , 1823) Leontopithecus rosalia (Linnaeus, 1766) Mico Lesson, 1840 Mico acariensis (van Roosmalen M.G.M., van Roosmalen T., M itterm eier et Rylands, 2000) Mi CO argent atus (Linnaeus, 1771) MicO chrysoleucos (Wagner, 1842) Mico emiliae (Thomas, 1920) Mico humeralifer (E . Geoffroy Saint-Hilaire, 1812) Mico intermedins (Hershkovitz, 1977) Mico leucippe Thomas, 1922 Mico manicorensis (van Roosm alen M .G .M ., van Roosmalen T., M itterm eier et Rylands, 2000) Mico marcai ( A lp erin , 19 9 3) Mico maiiesi ( M itterm eier, Schwarz et Ayres, 1992) Mico melanurus (E . Geoffroy Saint-Hilaire, 1812) Mico nigriceps (Ferrari et Lopes, 1992) Mico rondoni Ferrari, Sena, Schneider et Silva, 2010 Mico saterei ( Silva etNoronha, 1998) Oreonax Thomas, 1927 Oreonax flavicauda (Humboldt, 1 8 1 2 ) Pitheda Desmarest, 1804 Pithecia aequatorialis Hershkovitz, 198 7 Pithecia albicans Gray, i860 Pithecia ( irrorata ) hirsuta s p ix , 18 2 3 Pithecia ( irrorata ) irrorata Gray, 1842 Pithecia (irrorata) vanzolinii Hershkovitz, 1987 Pithecia ( monachus) milleri a lien . 1 9 1 4 Pithecia (monachus) monachus (E. Geoffroy Saint-Hilaire, 1812) Pithecia ( monachus) napensis Lonnberg, 1938 Pithecia ( pithecia) pithecia (Linnaeus, 1766) Pithecia (pithecia) chrysocephalai. Geoffroy Saint-Hilaire, 1850 Pithecia ( pithecia ) lotichiusi m ertens, 1925 Saguinus Hoffmannsegg, 1807 Saguinus bicolor ( s p ix , 1 8 2 3 ) Saguinus fuscicollis fuscicollis ( s p i x , 1 82 3) Saguinus fuscicollis avilapiresi Hershkovitz, 1966 Saguinus fuscicollis cruzlimai Hershkovitz, 1966 Table 1/2. References of scientific descriptions of all known Neotropical primates (present paper). 192 Marc G.M. van Roosmalen & Tomas van Roosmalen Sagliinus fuscicollis mura Rohe, Silva Jr., Sampaio et Rylands, 2009 Saguinus fuscicollis primitivus Hershkovitz, 1977 Saguinus fllSCUS (Lesson, 1840) Saguinus geoffroyi (Pucheran, 1845) Saguinus illigeri (Pucheran, 1845) Saguinus (imperator) imperator (Goeidi, (1907) Saguinus ( imperator) subgrisescens (L 6 n n b e rg , 19 40) Saguinus inustus (Schwarz, 1951) Saguinus labiatus labiatus (E. Geoffroy Saint-Hilaire, 1812) Saguinus labiatus rufiventer (Gray, 1 8 4 3) Saguinus labiatus thomasi (Goeidi, 1 9 o 7 ) Saguinus lagonotus (Jimenez de la Espada, 1870) Saguinus leucogenys (G ray, 1865) Saguinus leucopus (Gunther, 1876) Saguinus martinsi martinsi (Thom as, 1 9 1 2) Saguinus martinsi ochraceus h ershkovitz, 1966 Saguinus midas (Linnaeus, 1758) Saguinus my s tax my s tax (S p ix , 1 8 2 3) Saguinus mystax pileatus (I. Geoffroy Saint-Hilaire et D eville, 1848) Saguinus mystax plllto (L 6 n nb erg , 19 2 6) Saguinus niger (E. Geoffroy Saint-Hilaire, 1 803) Saguinus nig rifrons ( I. Geoffroy Saint Hilaire, 1850) Saguinus nigricollis nigricollis (S p ix , 1 82 3) Saguinus nigricollis graellsi ( Jimenez de la Espada, 1870) Saguinus nigricollis hernandezi h ershkovitz, 1982 Saguinus Oedipus (Linnaeus, 1758) Saguinus tripartitus (M iln e-E d w ard s, 1 8 7 8) Saguinus weddell i weddell i ( d e v ill e , 1849) Saguinus weddelli crandalli Hershkovitz, 1966 Saguinus weddelli melanoleucus (Miranda Ribeiro, 1912) Saimiri Voigt, 1 83 l Saimiri boliviensis boliviensis (I. Geoffroy Saint-Hilaire et de Blainville, 1 834) Saimiri boliviensis peruviensis Hershkovitz, 1984 Saimiri ( cassiquiarensis ) cassiquiarensis (Lesson, 1840) Saimiri ( cassiquiarensis ) albigena (von Pusch, 1942) Saimiri macrodon Elliot, 1907 Saimiri oerstedii oerstedii (Reinhardt, 1872) Saimiri oerstedii citrinellus (Thomas, 1904) Saimiri (SCilireuS) SCiureuS (Linnaeus, 1 758) Saimiri (sciureus) collinsi Osgood. 1 9 1 6 Saimiri UStUS (I. Geoffroy Saint-Hilaire, 1843) Saimiri vanzolinii Ayres, 19 8 5 Sapajus Kerr, 1792 Sapajus apella apella (Linnaeus, 175 8) Sapajus apella margaritae ( h oiiister, 1 9 1 4 ) Sapajus cay (iiiiger, 1 8 1 5 ) Sapajus flavins ( S c h re b e r, 17 7 4) Sapajus libidinosus ( s p ix , 1 82 3) Sapajus macrocephalus ( s p i x , 1 8 2 3) Sapajus nigritus ( G o l d f u s s , 1 8 09) Sapajus robustus (K uh l, 1 8 2 o ) Sapajus xanthosternos (Wied-Neuwied, 1826) Table 1/3. References of scientific descriptions of all known Neotropical primates (present paper). less intelligent, sensitive, and sociable than primates in general are - being forced to live as outcasts would equal a sure death. But, if it were healthy male individuals deviant from the socially selected skin and/or hair color pattern that are discriminated against merely for being slightly depilated or having its coat somewhat bleached somewhere, such young males pushed out of the group’s core area by high-ranking males will ally for the sake of survival alone. Their shared forced-upon marginal- ity could well drive them into looking beyond the horizon and together leaving the pack in search of a living space wherever it could be found. Once that living condition is fulfilled, they can start a new social group incorporating some females that they were able to attract from other resident groups on their way out. This phenomenon is known to com- monly take place in hierarchically structured primate societies that are ruled and defended by do- minant (alpha)-males (e.g., AlouattO) . It guarantees a certain primate to reach optimal densities in un- disturbed populations. Furthermore, it selects for males that are capable to lead and defend a social group. A 11-m ale parties of slightly eumelanin and/or pheomelanin bleached, or somewhat depilated males that are pushed out of their parental group’s living space and that follow the ‘trend to allopatry’, will range further and further away from the core of a taxon’s distribution. If suitable habitat to settle down is not encountered, the animals eventually will weaken, suffer from diseases, starve to death, or get predated upon. Very rarely, they happen to venture into for that species marginal or unsuitable habitat, being forced to adapt to an alien habitat or a different feeding niche. In extremely rare cases, such founder-groups or -colonies may diverge along this path into a different subspecies (whatever that may be) and eventually into a different species (whatever that may be) or ecospecies. This kind of sympatric speciation may have taken place in such cases as the cream-white, near-albinotic fair woolly monkey living year-round in the varzeas between On the origin of allopatric primate species 193 the lower Rio Javan and the right bank of the upper Rio Solimoes. Or, the pheomelanin bleached, over- all orange-colored woolly monkey from the head- waters of the Rio Jutai. Somewhat metachromic bleached founder-colonies of woollies driven by the ‘trend to allopatry’ once must have diverged from archetypic agouti-colored or saturated eumelanin ancestral La. poeppigU w hile adapting to a different ecological niche that was new to woolly monkeys - in this case that of a frugivorous, canopy-dw elling, brachiating inhabitant of white-water inundated floodplain forest (varzea). During our systematic surveys of primate distribution and diversity carried out in the matrix terra firme rain forest that stretches out behind the floodplain of some white-water rivers (e.g ., Javan, Jurua, Purus, Madeira), we were not able to detect any differences in phenotype between individual monkeys of a given taxon that we observed along the entire course (from source to headwaters) of each of these far-apart rivers. Contrary to what is the common presumption among prim atologists, this would mean that in territorial monkeys such as pygmy marmosets or saddle-back tamarins that occupy large distributions delineated by some of the largest tributaries of the Amazon, phenotypic characters of skin and pelage coloration, and/or local hair growth or depilation, seem to have stabilized across their entire distribu- tions. In other words, within the distribution of a given Amazonian monkey there does not exist something like a gradient of slightly different phen- otypes, color forms, morphs, or races. These obser- vations from the larger field have led us attributing full-species status to primate taxa like Cebuella pygmaea and C. niveiventris that are phenotypically stable throughout their (sometimes huge) distribu- tions. Consequently, we here introduced the concept of eco-species. This concept is firmly corroborated by the here proposed theory on the origin of allo- patric primate species. An ecospecies may be best defined as: “A genetically isolated population or group of populations of a kind that does not undergo gene flow from adjacent populations of one or more closely related kinds; and that shows a stabilized phenotype across the entire range in which it occupies a well-defined ecological niche, which it defends against any outside competitor, even beyond generic level:' This eco-species concept (ESC) avoids the often confusing arbitrary distinction between species, subspecies, race, morph, or form, for it adds sociobiological restric- tions to environmental (geographic, geomorpholo- gical and phy tosociological) ones that use to act on speciation and radiation in sociable territorial primates. Defined as such, the ESC may apply also to similarly socially structured mammals like coatis, peccaries, and some canids. In accordance to this definition, an enclave population of Callibella humilis that lives year-round in igapo forest fringing the Rio Atininga - genetically isolated from the main population that lives at least one hundred km to the north in primary terra firme rain forest - should be assigned a different species name in its own right. Or, in case the ranges of two Saddle-back Tamarins of the S. fuscicollis Clade, hitherto being treated as subspecies, are only separated by a nar- row contact zone - where its aggressive territorial defense effectively impedes any gene flow through cross-breeding or hybridization - each population should be given valid species status. But, wherever a former distributional boundary between two such ecospecies has been disrupted, removed by a vi- cariance, or overtaken by the more aggressive or opportunistic of two ecospecies, the latter will expand its distribution to the cost of the other. Then, a process of replacement is set in motion along a steadily moving frontline, which inevitably will lead to the extinction of the less aggressive, more vulnerable, or more sensitive of the two ecospecies. According to our doctrine of allopatric primate speciation this will always be the ecospecies that is the more advanced metachromic bleached one. Here we have mentioned at least four cases across the Amazon where such process of replacement (through physical extermination) of one primate by another is ongoing or about to be terminated: 1) the archetypic agouti, gray, and dark red-brown coated Lake Baptista Titi Monkey Callicebus baptista extending its range along the southbank of the Rio Amazonas to the cost of the advanced bleached, yellow- and-gray coated Hoffmanns’s Titi Monkey Callicebus hoffmannsi and the near- a lb in otic eco- species from the right bank of the Rio Mamuru; 2) the archetypic saturated eumelanin Midas Tamarin Saguinus midasvz rsus the progressively bleached, halfway to fully albinotic Pied Two-colored Tam- arin S. bicolor (including S. OchraceUS) and Mar- tins’s B are-face Tamarin S. tnartinsi , the latter three ecospecies being currently at the verge of extinction caused by a rapid southern expansion of midas 194 Marc G.M. van Roosmalen & Tomas van Roosmalen (Fig. 10); 3) the saturated eumelanin Weddell’s Saddle-back Tamarin S. (fusticollis ) Weddelli ex- panding its range to the cost of the near-albinotic Rondon’s Marmoset MlCO rondoni, pushing the frontline eastward into the interfluve delineated by the Rios Guapore and Ji- Parana after having traversed the upper Rio Madeira in the recent past; 4) Gray’s Saki Pithecia hirsuta (or P. mittermeieri) extending its range northwards to the cost of the near-albinotic B u ffy Saki taxon P. CllbicCMS. In cases of replacement it is always the more advanced metachromic bleached to albinotic ecospecies that is loosing the battle and eventually will go extinct. Though only documented by us in semi-captive and free-ranging, but artificially composed multi- pecies populations, during social conflicts it was invari- ably the more advanced metachromic bleached in- dividual monkey or group of monkeys that suffered from dominant-male discriminatory behavior, being bullied, repeatedly physically attacked or violently assaulted, and eventually forced out of the core (compound) area, where we provided additional food on feeding platforms constructed up in the canopy. If not moving voluntarily to the periphery, so turning into outcasts, these monkeys could be bitten to death by the invariably less bleached, more aggressive, conspecific leading male(s). In retro- spect, we recall that all neotonic, advanced meta- chromic bleached and near-albinotic individual monkeys kept free-ranging in our respective halfway-houses by comparison were invariably more soft-hearted, more sensitive, cooperative, adaptable, and (not surprisingly?) smarter than the male congeners by whom they were discriminated, pushed into the periphery, or banned from the core area. Applying these observations to the wild, the trend to allopatry boosted by seemingly non-ad- aptive social selection - leading males that discrim- inate upon phenotypically deviant mutant young males - in evolutionary sense could well turn out to be truly adaptive. To cite Charles Darwin (1 859): “ In the long history of humankind -and animalkind, too- those who learned to collaborate and impro- vise most effectively have prevailed ’ .And: “ It is not the strongest of the species that survives, or the most intelligent that survives. It is the one that is the most adaptable to changer Applying the doctrine to the evolution of hom- inins, in particular Homo sapiens, one may ponder and speculate about questions like the following: “ Why, and driven by what force about six million years ago somewhere in Tropical Africa an ape-like lineage of primates -our hominid ancestors- left the rain-forest canopy and ventured into an arid open- savanna scrub landscape ? ” The common ancestors of the Great Apes and the human line of hominins ( Homo ) were arboreal primates that had adopted brachiation (suspended arm -over-arm -sw inging underneath the twig/branch substrate) as a special locomotor pattern. Brachi- ation allows large-bodied arboreal primates to quickly move through the canopy and get to the fleshy fruits that are, as is the rule in any tropical rain-forest environment, distributed in the far peri- phery (small- branch/twig micro-habitat) of canopy- and emergent-tree tops. Brachiation is a primarily arboreal type of locomotion that evolved exclus- ively in some Neotropical Monkeys (i.e., spider, woolly and woolly spider monkeys) as well as in the Old-World Apes (i.e., gibbons, siamangs, bonobos/pygm y chimps, chimpanzees, orang-utans, and gorillas). It may never have evolved in Prosimi- ans, which are the more primitive among all the world’s primates. It followed an independent evol- utionary path, a convergent or parallel evolution, in a physiognom ically similar natural environment - the tropical forests of Southeast Asia, Central Africa, and South America (the larger Amazon Basin). A major intercontinental difference is that some monkeys in the Neotropics developed a pre- hensile tail as extra support in suspensory loco- motion, therefore called “sem i-brachiation”, whereas apes during the evolutionary process toward brachiation lost a functional tail. Brachiation without use of a fifth limb is called “true brachi- ation”. Most plausibly, our early ape-like hominid ancestors that about 6 M YA descended from the can- opy of C entral-A frican rain forest much resembled extant Spider Monkeys in their general locomotor pattern and diet. Brachiation is associated with a dietary preference for ripe, pulpy, nutritious fruits that contain a single to few large seeds. The upright position of the trunk associated with an arboreal life-style involving much brachiation happened to be a crucial pre-adaptation for later bipedal (two- legged) upright walking on the ground. It enabled our early ancestors to leave the trees in the same way as gorillas once did, but different in that the Great Apes adopted ‘knuckle-walking’ as the principal locomotor pattern to walk on the ground. On the origin of allopatric primate species 195 Similarities between Spider Monkeys and Chim- panzees are striking as we consider that at least twenty-five million years of evolution on different continents do separate these primates from one an- other. Cognitive features that both brachiating primates share are the mental capacity to visualize, pre-plan, and map out in time and space complex economic foraging routes to be followed that very day, tomorrow, the day after tomorrow, and perhaps even over several days ahead. Moreover, these primates are able to lay out these foraging routes across a landscape that is covered with dense trop- ical rain forest containing only few seasonal, widely dispersed food sources at any given time (Van Roosmalen, 1 985a; 2013a). Consequently, both spider monkeys and pygmy chimpanzees (bonobos) may well depict a marked period or stage in the evolution of our early ancestors that may have specialized first in feeding upon ripe, juicy, lip id - and protein-rich, large-seeded fruits. Perhaps, that feeding niche may have been the condition that predestined our ancestors, both locomotorily and mentally, to leave the trees and become two-legged ground-dwelling foragers with an advanced use of the hands (e.g ., dexterity, precision grips, tool- fashioning). And at the same time growing big babies and three to four times bigger brains (Lynch & Granger, 2008). In physical, anatomical, physiolo- gical, and mental respect, therefore, descending from the trees and adapting locomotorily to bipedal walking and running over the ground was not the ‘near-impossible’ step that it may seem to be. If we put it in Darwinian evolutionary perspective, how- ever, to let it happen, until now an intraspecific social driver was missing that must have acted on the undoubtedly territorially and hierarchically organized communities of these ape-like ancestors with the brain size of contemporary chimpanzees (400 cc). Forthcoming our thirty-five years living in the Amazon and conducting long-term research on captive, feral, as well as wild monkeys - the latter mostly representing pristine populations that were never in any way disturbed by humans - we here suggest the ‘trend to allopatry’ among slightly depilated and/or metachromic bleached male indi- viduals (mutants) in primate populations being the principal force that has driven founder-colonies of our early ancestors - for the mere sake of survival - out of their preferred habitat -canopy trees- into (to them) new, with respect to natural enemies risky and hostile landscapes. A s sociable and intelligent mammals suffering from intraspecific population pressures and discriminatory social constraints, outcast males must have taken on the challenge to traverse whatever barrier on their way out. So, they ventured into the arid, in many aspects hostile natural environment of savanna scrub and open woodlands. In a similar way as a small population of Gracile Capuchins on the slopes of tepuis like Pico da Neblina successfully adapted to a predom - inantly ground-dwelling life-style; the Mountain Gorilla successfully adapted to a fully terrestrial life-style in the cloud forests of the Virunga vulca- noes in C entral A fric a ; the Western Chimpanzee of the ‘subspecies’ VerilS once adapted to a predom- inantly terrestrial life-style in an arid, for specialist frugivores inappropriate or marginal natural envir- onment - the open savanna scrub of West Africa (Patterson et al., 2006); the near-albinotic Rio Javari Fair Woolly Monkey with an overall cream-white colored coat, and the Rio Jutai Orange Woolly Monkey with an overall orange colored coat, ad- apted to varzea floodplain forest along the upper Amazon and lower Javari Rivers, and the upper Jutai River, respectively; the Peruvian Yellow -tailed Woolly Monkey in complete isolation adapted to high-altitude cloud forest in the NE Peruvian Andes; the advanced pheomelanin to near-albinotic Bald-headed U akaris adapted to seasonally inund- ated white-water floodplain forest (varzea) along the Amazon River and some of its southern tributa- ries that drain the southeastern flanks of the Andes; among others. Looking at the distribution of C en tral- A m eric an spider monkeys of the Ateles geojfroyi Clade, we could speculate about an imaginary evolutionary path that could have been followed by an advanced metachromic bleached, near-albinotic founder- colony of the Central Amer- ican Yucatan Spider Monkey Ateles ( geojfroyi ) yucatanensis from the tropical forest of Yucatan Peninsula in SE Mexico. By the ‘trend to allopatry’ forced out of the canopy of a semi-deciduous rain forest somewhere on the Yucatan Peninsula - the taxon’s current deadend distribution - some founder- colony may venture into the savanna and desert scrub of SE Mexico and from there further into the Midwest of the US. To survive in such (for spider monkeys) alien landscape it would quickly have to loose a functional tail and adopt bipedal upright walking as its main locomotor pattern. It is tempting 196 Marc G.M. van Roosmalen & Tomas van Roosmalen to imagine a similar scenario for progressively metachromic bleached, depilated, red- or white- skinned near-albinotic early hominids 6 MYA radiating away from their archetypic, saturated- eu melanin congeners they had in common with ancestral chimpanzees. Driven by the trend to allo- patry, in a similar way founder-colonies may have left the semi-deciduous rain forests of C Africa and ventured first into the savannas, plains and desert scrub of N Africa and, thereafter, into the tem- perate-clime dominated landscape of S + C Europe, the Middle East and SE Asia. Recent evidence from molecular biology suggests that it took several hun- dreds of thousands years for our early ancestors to evolve in two distinct animals: the open savanna explorers leading toward proto-humans, and those remaining arboreal resulting in chimpanzees (Pat- terson et al., 2006). In accordance with recent phylogenetic research, the modern Chimpanzee Pan troglodytes diverged from the proto- or ar- chetypic, saturated eumelanin, overall blackish- brown colored Bonobo (Pygmy Chimpanzee) Pan panisCUS . The common Chimpanzee is an oppor- tunist having an omnivorous diet, whereas the Bonobo holds a predominantly specialist frugivor- ous diet. In comparison to common Chimpanzees, Bonobos are egalitarian, peaceable, non-violent creatures that live in loosely organized, matriarchal social groups in which the males may defend their territories, but rather adopt a “M ake Love No War” philosophy of life. Bonobos have never been repor- ted to involve in raids on neighboring group males, whereas common chimpanzees have been seen performing a kind of troop-hunting culture in which beta-males led by one alpha-male sometimes do attack neighboring males or small mixed parties, killing and eating some of them. Bonobos live in the dense tropical rain forests of Central Congo. Their distribution is thought to represent the cradle of chimpanzee evolution or the center of chimpan- zee (genus Pan) dispersion. Applying Hershkovitz’ hypothesis of m etachrom ism , Chimpanzees may well have derived from (proto)-B onobos. Nowadays, the two species are allopatric. The trend to allopatry may have forced ancestral chimpanzees to swim across or circumvent the Congo River that does act as a geographic barrier in present-day distributions. The farther in any but southern direction from the center of Pan troglodytes dispersion, located just north of the Congo River, the more arid the landscape becomes, the more often chimps do descend from the trees and ‘knuckle-walk’ on the ground, and the more chimps have adapted to what bonobos would consider inappropriate or marginal habitat - unsuitable to highly specialized mature fruit-eaters that bonobos are. At the same time, we see chimpanzees becoming more pheomelanin to euchromic bleached, their skin getting lighter colored (less pigmented), their coat thinner and locally depilated or almost hairless, and elderly individuals becoming gray with age. Another question to ponder about with the doc- trine in mind: “ Why, and driven by what force some of our Homo ancestors between 100,000 and 50.000 years ago left the origin and center ofhom- inid dispersion -Central and North Africa being considered the cradle of human evolution- to ven- ture into the clime- and habitat-wise new, but un- suitable or (at least ) marginal landscape of Europe, the Middle East and Asia?” A fter Homo ereCtUS having grown much bigger brains on the plains, some millions of years later the trend to allopatry may have been again the principal driving force for some founder-colonies of Homo Sapiens to move ‘Out of A frica’ . The pioneers that ventured into new landscapes to the north could do so only by occupying an ecological feeding niche that was new to former small-game hunter-fisher- gatherers, that of big-gam e hunter-gatherers. H ere by, the invention to first carrying along fire, soon fol- lowed by the skill to kindle it, was essential in the adaptation process to a new feeding niche, as their (our) digestive system is not apt to decompose raw meat. It has to be cooked or barbecued. Apparently, in very low densities - recent estimates place the population of Europe 30,000 years ago at about 5.000 people - these humans following herds of prehistoric megafauna (e.g., mammoth) and driving them to extinction in the Holocene, have spread rapidly across the whole of Europe and Southeast Asia, one route taking them as far as Australia and Tasmania, the other to the far northeastern corner of Siberia. From these places they eventually could reach and inhabit some Pacific Islands, and most amazingly also the continent of South America, first about 30-40,000 years ago by bordering the Antarc- tic during one of the glacials, and a second time, about 15,000 years ago, via Beringia and North America (Van Roosmalen, 2013c). We could ask ourselves if these all could have been advanced On the origin of allopatric primate species 197 metachromic bleached, euchromic to albinotic founder-colonies or colonizing parties that were pushed out from dead-end distributions in Africa and Asia following the male-territorial primate-born trend to allopatry? ACKNOWLEDGMENTS We would like to dedicate the theory laid down in this paper to the memory of Alfred Russel Wal- lace (1 823-19 1 3), who inspired us to follow his early footsteps into the Amazonian realm. 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