Early Palaeogene Louisinidae (Macroscelidea,Mammalia), their relationships and north European diversity

Seventeen species from the Palaeocene and Early Eocene of northern Europe, of which 12 are new, are described belonging to the extinct macroscelidean family Louisinidae, raised here from subfamily rank. These species belong to nine genera, of which five are new. The new genera are Walbeckodon , Berrulestes , Gigarton , Thryptodon , and Prolouisina . The new species are Walbeckodon krumbiegeli , Walbeckodon girardi , Paschatherium levei , Berrulestes phelizoni , Berrulestes pellouini , Berrulestes poirieri , Gigarton meyeri , Gigarton sigogneauae , Gigarton louisi , Thryptodon brailloni , Louisina marci , and Teilhardimys brisswalteri . Prolouisina is erected for ‘Louisina’atavella Russell, 1964. Cladistic analysis was undertaken to understand the relationships within the Louisinidae and between them and the North American family Apheliscidae, in which they had earlier been included as a subfamily. Louisinidae are shown to be sister group to a clade consisting of Apheliscidae plus Amphilemuridae and part of a paraphyletic and polyphyletic Adapisoricidae, all of which are tentatively considered to be stem members of the order Macroscelidea. The most primitive macroscelidid, Chambius, from the Early Eocene of northern Africa is nested within Apheliscidae when postcranial characters were included, but in a majority of cases within the Louisinidae when postcranial characters were excluded. Most species from northern Europe became extinct at the end of the Palaeocene, although the genus Paschatherium survived for much of the Early Eocene and Teilhardimys survived into the earliest Eocene. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 856–936.     

Wasatchian (Early Eocene) pollen floras from the Red Hot Truck Stop, Mississippi, USA

Sediments at the Red Hot Truck Stop (RHTS), Mississippi, USA are important because they contain the lowest latitude record of both the earliest known Eocene plant and mammal fossils in North America. The RHTS contains the uppermost Tuscahoma Formation and the lowermost part of the basal Bashi and Hatchetigbee formations. The Tuscahoma Formation is composed of glauconitic sands and silts that represent estuarine to shallow marine sediments. Faunal remains indicate that the RHTS section belongs to the Wasatchian North American land mammal age and specifically to the lower Graybullian subdivision. Pollen and spore floras from the RHTS are moderately diverse (113 taxa) and contain families that today are associated with warm‐adapted vegetation types such as Annonaceae, Bombacaceae, Burseraceae and palms. Eocene first occurrences are represented by Brosipollis sp., Celtispollenites sp., Interpollis microsupplingensis, cf. Nuxpollenites psilatus, Platycarya spp., Retistephanocolporites sp. and Symplocos? contracta and by one genus of pteridophyte spore (Granulatisporites sp.). The overall composition and within‐sample diversity of the sporomorph flora is more similar to the Hatchetigbee Formation (early Eocene) than to the middle Tuscahoma Formation (late Palaeocene) but among‐sample diversity remains unchanged across the Palaeocene/Eocene boundary. The distinct composition of the RHTS demonstrates that floral change across the Palaeocene/Eocene boundary is complex and composed of several phases of floral change.     

New proviverrines (Hyaenodontida) from the early Eocene of Europe; phylogeny and ecological evolution of the Proviverrinae

We describe six proviverrine species from the Early Eocene of France. Three species are new: M inimovellentodon russelli sp. nov. from Mutigny [mammal palaeogene (MP)8 + 9], B oritia duffaudi sp. nov. from La Borie (MP8 + 9), and L eonhardtina godinoti sp. nov. from Grauves (MP10). We describe new specimens and propose new generic combinations for three species from MP10: Protoproviverra palaeonictides, Matthodon menui, and Oxyaenoides lindgreni. We also propose a new generic combination for the primitive Eoproviverra eisenmanni (MP7). Matthodon menui was previously considered as a possible oxyaenodontan, but the new fossils clearly support its reference to Hyaenodontida. Leonhardtina godinoti and Ma. menui are the oldest occurrences for these genera, which were previously unknown before the Middle Eocene. Moreover, the discovery of the proviverrine Mi. russelli in Mutigny implies that the Proviverrinae dispersed in Northern Europe between biozone Palaeocene‐Eocene (PE) III (Abbey Wood) and biozone PE IV (Mutigny). This also supports a homogenization of the European faunas during the Early Eocene. The dispersal is concomitant with the disappearance of the oxyaenodontans, arfiines, and sinopines (Hyaenodontida) from Europe. The proviverrines may have filled the ecological niches left vacant by the disappearance of the other carnivorous mammals. With 20 genera and over 30 species, proviverrines were successful in Europe. We performed the first phylogenetic analysis comprising almost all the Proviverrinae. Our analyses indicate that the Proviverrinae diversified greatly during the Early Eocene Climatic Optimum and show a general trend towards specialization throughout the Eocene. © 2014 The Linnean Society of London     

Taxonomic and biostratigraphical re-assessments of Subterraniphyllum Elliott (Corallinales, Rhodophyta)

A taxonomic and biostratigraphical re‐assessment of Subterraniphyllum Elliott (Corallinales, Rhodophyta) is presented. Results from studies of the type collection and of newly collected material from north‐eastern Italy and northern Slovenia have shown that this taxon is not a geniculate coralline red alga as originally suggested by Elliott and most subsequent authors. Vegetatively, Subterraniphyllum most closely resembles certain living members of the Corallinales; however, the phenetic and phylogenetic relationships of Subterraniphyllum to other Corallinales cannot be determined with greater certainty. The exclusion of Subterraniphyllum from any group of Corallinaceae with genicula is based on unequivocal evidence that branch formation does not involve the occurrence of genicula. Subterraniphyllum seems to be restricted to the Oligocene. Reports of occurrences in Upper Eocene and Lower Miocene sediments cannot be substantiated. Subterraniphyllum, however, cannot be considered a useful stratigraphical marker until further data on its occurrence in well‐dated carbonate sequences are acquired. This study illustrates the problems associated with placing fossil coralline algal specimens into geniculate genera without the preservation of relevant morphological characters. This is especially true in the absence of the careful assessment of fossil material with respect to current taxonomic concepts of geniculate coralline genera, all of which are based on studies of living species. According to the current concepts for geniculate coralline genera, the placing of fossil specimens into geniculate genera without appropriate evidence must be avoided by grouping all potentially geniculate fragments under the informal group ‘Geniculate sensu lato’. Furthermore, for all those many fossil specimens where unequivocal evidence is not present, it is possible to utilize ‘form genera’ based on characters that are normally preserved. This leads to creating a consistent, workable system of applying names to most fossil corallines so that they can be reliably used in relation to stratigraphical and palaeoecological studies.     

Thanetian–Ilerdian coralline algae and benthic foraminifera from northeast India: microfacies analysis and new insights into the Tethyan perspective

Thanetian–Ilerdian carbonate deposits from the Lakadong Limestone in Assam Shelf, Meghalaya, northeast India, are studied with respect to microfacies distributions and controlling ecological factors on dominant biogenic components. Palaeoenvironmental implications are inferred following the detailed analysis of microfacies characterized by rich assemblages of coralline red algae and benthic foraminifera. The carbonate sediments have been interpreted as lagoonal to outer shelf facies. It is envisaged that the analysed benthic communities thrived in a meso‐oligotrophic regime above the fair‐weather wave base. The Lakadong Limestone constitutes a well‐preserved record of Late Palaeocene–earliest Eocene shallow marine carbonate ecosystem and has high potential to decipher its response to an interval of distinct changes in climate and tectonic settings. The abundance of oligotrophic larger benthic foraminifera in the Lakadong Limestone is comparable to the foraminiferal assemblages of west Tethys. The phylogenetic changes (‘Larger Foraminiferal Turnover’, LFT) and subsequent rapid radiation of typical Eocene larger benthic foraminifera (Alveolina, Nummulites) usually observed in the west Tethys have also been observed in the upper part of the Lakadong Limestone. The eastward migration of Eocene foraminifera from the west coincided with the India‐Asia collision and global warming events at the Palaeocene–Eocene boundary that may have produced a wide array of modifications in biogeography, seasonal run‐offs and ocean circulation pathways. The data indicate that rapid rate of migration from west before the onset of geographic barriers and/or timely restoration of pan‐Tethyan environmental conditions ensured the incidence of these forms in the earliest Eocene sediments.     

A new taeniolabidoid multituberculate (Mammalia) from the middle Puercan of the Nacimiento Formation,New Mexico,and a revision of taeniolabidoid systematics and phylogeny

Multituberculates were amongst the most abundant and taxonomically diverse mammals of the late Mesozoic and the Palaeocene, reaching their zenith in diversity and body size in the Palaeocene. Taeniolabidoidea, the topic of this paper, includes the largest known multituberculates, which possess highly complex cheek teeth adapted for herbivory. A new specimen from the early Palaeocene (middle Puercan; biochron Pu2) of the Nacimiento Formation, New Mexico represents a new large‐bodied taeniolabidoid genus and species, K imbetopsalis simmonsae . A phylogenetic analysis to examine the relationships within Taeniolabidoidea that includes new information from K imbetopsalis gen. et sp. nov. and gen. nov. and from new specimens of Catopsalis fissidens, first described here, and data from all other described North American and Asian taeniolabidoids. This analysis indicates that Catopsalis is nonmonophyletic and justifies our transfer of the basal‐most taeniolabidoid ‘Catopsalis’ joyneri to a new genus, V alenopsalis . K imbetopsalis and Taeniolabis form a clade (Taeniolabididae), as do the Asian Lambdopsalis, Sphenopsalis, and possibly also Prionessus (Lambdopsalidae). Taeniolabidoids underwent a modest taxonomic radiation during the early Palaeocene of North America and underwent a dramatic increase in body size, with Taeniolabis taoensis possibly exceeding 100 kg. Taeniolabidoids appear to have gone extinct in North America by the late Palaeocene but the appearance of lambdopsalids in the late Palaeocene of Asia suggests that they dispersed from North America in the early to middle Palaeocene.     

Cenozoic deep-sea ostracods from Maud Rise, Weddell Sea, Antarctica (ODP Site 689): a palaeoceanographical perspective

Cenozoic palaeoceanography of the Maude Rise, Weddell Sea, Antarctica, has been investigated using Palaeocene to Quaternary deep-sea ostracod faunas from 23 samples of ODP Site 689. The abundance of ostracods is high enough only during the Palaeogene (Palaeocene-Oligocene) to allow palaeoceanographical inferences based on changes in diversity, dominance, endemism and faunal turnover (first and last occurrences). The abundance is particularly high throughout the Palaeocene and Eocene, but declines irreversibly near the Eocene/Oligocene boundary. The diversity increases more or less continuously from the Early Palaeocene to the Middle Eocene, and then it generally decreases throughout the remaining part of the Palaeogene (Middle Eocene-Oligocene); an exception is a positive peak in the Shannon-Weaver index in a single sample in the Late Oligocene. No positive peaks in diversity and taxa originations (first occurrences) at c. 40-38 Ma, occurs at Site 689; so the site provides no evidence for the establishment of the psychrosphere at this time. This corroborates similar regional results from an earlier study of benthonic foraminifera. Explanations for this may be related to Late Eocene-Early Oligocene changes in sedimentology and clay-mineralogy (associated with the progressive cooling of the Antarctica) which could have negatively affected abundance and diversity locally at Site 689. Alternatively, by this time, the ostracod fauna could also have been subjected to selective removal (with possible local extinction) of taxa (due to increased ventilation) or to thanatocoenosis dissolution (due to a decrease in temperature and availability of CaCO₃). A further possibility may be related to the fact that Site 689 was at intermediate water depths and may have remained within older water masses near the Eocene/Oligocene boundary. Failing these explanations, the results could indicate that the Late Eocene-Early Oligocene palaeoenvironmental changes in the world oceans were more gradual and occurred over a longer time interval than the global ostracod data show, at least at southern high latitudes.     

A highly derived anomalurid rodent (Mammalia) from the earliest Late Eocene of Egypt

Abstract: A new genus and species of diminutive anomalurid rodent, Shazurus minutus, is described on the basis of 15 isolated teeth from the earliest late Eocene (approximately 37 Ma) Birket Qarun Locality 2 in the Fayum Depression of northern Egypt. Shazurus is surprisingly specialized for its age, being most similar in dental morphology to early Miocene Paranomalurus and extant Anomalurus, and is quite different from the roughly contemporaneous anomaluroid genera Nementchamys and Pondaungimys from Algeria and Myanmar, respectively. Parsimony analysis of dental features places Shazurus within crown Anomaluridae as a sister group of two species of Paranomalurus to the exclusion of extant Anomalurus and Idiurus. The marked morphological differences between the two oldest Afro‐Arabian anomaluroids (Shazurus and Nementchamys) suggest that the taxa share a much more ancient common Afro‐Arabian ancestor, possibly derived from Zegdoumyidae. Isolated teeth of Shazurus and other Eocene anomaluroids reveal little about their palaeobiology, but the complete absence of Anomaluroidea from the younger (late Eocene to early Oligocene) Jebel Qatrani Formation is presumably a reflection of environmental change through the late Eocene in northern Africa.     

Palaeocene origin of the Neotropical lineage of cleptoparasitic bees Ericrocidini‐Rhathymini (Hymenoptera,Apidae)

Cleptoparasitic bees abandoned pollen‐collecting for their offspring and lay their eggs in other bees' provisioned nests. Also known as cuckoo bees, they belong to several lineages, but are especially diverse in Apinae. We focused on a lineage of apine cleptoparasitic bees, the clade Ericrocidini + Rhathymini, which attack nests of oil‐collecting bees. We sequenced five genes for 20 species in all genera of this clade plus a large outgroup to reconstruct the phylogeny and estimate divergence times. We confirmed the monophyly of the clade Ericrocidini + Rhathymini and its position inside the ericrocidine line together with the tribes Protepeolini, Isepeolini, Osirini and Coelioxoidini. Our results corroborated the current taxonomic classification. Ericrocis is sister to all Ericrocidini and the position of Acanthopus and the most diverse genus Mesoplia were inconclusive. Ericrocidini + Rhathymini diverged from other apine cleptoparasitic lineages 74 Ma in the Cretaceous and from each other in the Palaeocene at 61 Ma. Considering the robust molecular evidence of their sister relationships, the striking differences in the morphology of first‐instar larvae of the two groups may represent adaptations to the nesting biology of their hosts. As other cleptoparasites in the ericrocidine line, Ericrocidini and Rhathymini possess larvae which are adapted to kill the immature host and to feed on floral oil provided by the host female. The evolution of host specialization in the line Ericrocidini + Rhathymini retroceded to the Eocene when they differentiated synchronously with their hosts, Centris and Epicharis .     

Taxonomic review of the “Bulimes”, terrestrial gastropods from the continental Eocene of the Hamada de Méridja (northwestern Sahara,Algeria) (Mollusca: Stylommatophora: Strophocheilidae?), with a discussion of the genera of the family Vidaliellidae

Terrestrial gastropods occur in many North African localities in Eocene continental deposits. Here we analyse the faunal assemblage from the Hamada de Méridja Formation in southwestern Algeria, dated as Early to Middle Eocene on the basis of charophytes. The assemblage consists of three closely related species that to date have been classified either in the extant Madagascan genus Leucotaenius v. Martens, 1860, or in the SW European Eocene genera Romanella Jodot, 1957 and Vicentinia Jodot, 1957. This is rejected for shell morphological and phylogeographical reasons, and a new classification as Maghrebiola gen. nov. is proposed. Maghrebiola is tentatively placed in the South American family Strophocheilidae, as species from the Early Eocene Itaboraí Basin of Brazil, currently placed in the genus Eoborus Klappenbach and Olazarri, 1970 in the family Strophocheilidae, superfamily Acavoidea, have a very similar shell habitus. This record possibly extends the known geographical range of the Strophocheilidae into the African continent during the Eocene. Immigration of this stock into North Africa during the Cretaceous via a still existing plate connection is assumed. An attribution of Maghrebiola to the African family Achatinidae is unlikely for shell morphological reasons despite certain habitus similarities, although the Priabonian genera Arabicolaria and Pacaudiella from Oman most likely belong into this family, and not to the Vidaliellidae as originally proposed. Possible causes for the very low diversity of the assemblage are mainly unfavourable living conditions, i.e. a relatively dry climate resulting in sparse vegetation and only occasional presence of water bodies, which may have had increased salinities, accounting for the lack of freshwater mollusks. The absence of any competing large gastropods may possibly have facilitated high intraspecific variability leading to sympatric occurrence of three closely related species, due to the animals occupying a wide range of available ecological niches. As the species discussed here have also been attributed to the genera Romanella and Vicentinia in the Vidaliellidae, we provide an appendix with annotated characterisations of most genera of the Vidaliellidae and list the nominal species assigned to them. This family is tentatively placed in the South American superfamily Orthalicoidea; its stock would have similarly immigrated from South America, but have successfully colonized mainly SW Europe, with only one Eocene species [Romanella kantarensis (Jodot, 1936)] recognized in Algeria.     

First record of Miocene crustaceans from Hormozgan Province,Southern Iran

Four decapod crustacean species from the Middle Miocene Mishan Formation of Hormozgan Province, Southern Iran are described. Three species of leucosiid crabs represent the first fossil record for their respective genera outside the Indo-Pacific region. The oldest records for Leucosia sensu lato, Phylira sensu lato, and Arcania are from Middle Miocene deposits of the Indo-Pacific, and thus their paleobiogeographic distribution is extended for this lapse of time. Harpactocarcinus miocenicus n. sp. represents the youngest and most oriental occurrence for a typically Eocene genus, whose species are known mainly from Central Europe. This contribution represents the first formal report for Tertiary crustaceans from Iran.     

The long‐term history of dispersal among lizards in the early Eocene: new evidence from a microvertebrate assemblage in the Bighorn Basin of Wyoming,USA

Abstract: Early Eocene mammal faunas of North America were transformed by intercontinental dispersal at the Paleocene–Eocene boundary, but lizard faunas from the earliest Eocene of the same area were dominated by immigrants from within the continent. A new lizard assemblage from the middle early Eocene of Wyoming sheds light on the longer‐term history of dispersal in relation to climate change. The assemblage consists of three iguanid species (including two new species possibly closely related to living Anolis), Scincoideus, ‘Palaeoxantusia’, four anguids, two species of an undescribed new anguimorph clade, Provaranosaurus and a varanoid (cf. Saniwa). Most North American glyptosaurin glyptosaurines are now referred to Glyptosaurus, and Glyptosaurus hillsi is given a new diagnosis. Scincoideus is otherwise known only from the mid‐Paleocene of Belgium, and the specimens described here are the first to document intercontinental dispersal to North America among lizards in the early Eocene. Like in mammals, some immigrant lizard lineages first appearing in the Bighorn Basin in the earliest Eocene persisted in the area long after the Paleocene–Eocene thermal maximum, but other immigrants appear to have been restricted to the Paleocene–Eocene thermal maximum.     

Mycoflora of camel and goat hairs from Al-Arish,Egypt

The frequency of occurrence of fungi in 120 hair samples of camel and goat from four different localities of Al-Arish governorate was determined.Twenty-six genera and 54 species were collected from the two substrates and the most common genera were Chrysosporium and Aspergillus, followed by Cladosporium. From the preceding genera Chrysosporium keratinophilum, C. tropicum, C. indicum, Aspergillus flavus, A. niger and Cladosporium cladosporioides. Also, other keratinophilic fungi were isolated such as C. luteum, C. pannorum, C. parvum, C. dermatitidis, C. inops, Arthroderma tuberculatum, Histoplasma capsulatum and Myceliophthora vellerea.     

Variation in the shape of the palate in adapis (Eocene,Adapiformes) compared with living primates

This study is a quantitative analysis of the variation of palate shape inAdapis, an Eocene fossil primate. The observed coefficients of variation of palate dimensions inAdapis are statistically greater than in most living primate species. The histogram of the index (palate width/palate length) emphasizes the great heterogeneity in the fossil species. Two allometric scalings of palate dimensions (length and width) on condylobasal length were performed. Plots of perpendicular distances from the two regression lines show that living species are always more concentrated around the middle of the diagrams than isAdapis. The space occupied byAdapis is as great as, or even greater, than those occupied by living genera. Hence, several species are certainly presenent in the fossil sample, and the varieties distinguished by Stehlin in 1912, are in fact valid species names.     

Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae)

Most of the 29 living species of Potamididae show a close association with mangroves. The trees provide the snails with shelter, protection from predators, a solid substrate and sometimes food. Using sequences from three genes (nuclear 18S rRNA and 28S rRNA, mitochondrial COI) we derive a molecular phylogeny and recognize six living genera (Terebralia, Telescopium, Tympanotonos, Cerithidea, Cerithideopsis, Cerithideopsilla). The oldest modern genera (Terebralia, Cerithideopsis) appeared in the Tethyan realm in the Middle Eocene, shortly after the origin of mangrove trees. Whereas most potamidid genera are now restricted to either the Indo-West Pacific (IWP) or to the eastern Pacific plus Atlantic (EPA), sister clades of Cerithideopsis survive in both realms. Based on a reinterpretation of the fossil record (particularly of the monotypic Tympanotonos and extinct Potamides), and parsimonious reconstruction of ancestral habitats, we suggest that the living potamidids are an adaptive radiation that has always been closely associated with mangroves. The specialized tree-climbing groups Cerithidea and Cerithideopsis were independently derived from mud-dwelling ancestors. Cerithideopsilla cingulata (a species complex in the IWP) and 'Potamides' conicus (in the Mediterranean and Indian Ocean) form a single clade within the genus Cerithideopsilla. This refutes the hypothesis that 'P.'conicus is the sole relict of the Tethyan Potamides that has occurred in the Mediterranean region since the Palaeocene. Instead, the phylogeny and fossil record suggest that an ancestor of Cerithideopsilla conica with planktotrophic larvae dispersed from the IWP to the Mediterranean in the Middle Miocene, that its direct development evolved in the Mediterranean during the Pliocene, and that it reinvaded the Indian Ocean during the Plio-Pleistocene.     

Tortoise (Reptilia,Testudinidae) radiations in Southern Africa from the Eocene to the present

Africa, inclusive of the West Indian Ocean islands, harbours 11 of the world's 16 extant testudinid genera. Fossil records indicate that testudinids originated in Asia and dispersed first to North America and Europe (Early Eocene) and later to Africa (Late Eocene). We used mitochondrial (1870 bp) and nuclear (1416 bp) DNA sequence data to assess whether molecular data support the late cladogenesis of Southern African testudinid lineages. Our results revealed strong support for the monophyly of a clade consisting of Kinixys, the two Malagasy genera and four Southern African genera (Psammobates, Stigmochelys, Homopus and Chersina). Kinixys diverged from this clade in the Late Palaeocene, suggesting that testudinids occupied Africa at an earlier date than indicated by fossil records. The Southern African tortoises consist of three, strongly supported clades: Psammobates + Stigmochelys; the five‐toed Homopus + Chersina; and the four‐toed Homopus. Due to the paraphyly of Homopus, we propose the taxonomic resurrection of Chersobius for the five‐toed Homopus species (boulengeri, signatus and solus). Cladogenesis at the genus level occurred mainly in the Eocene, with Chersina and Chersobius diverging in the Oligocene. The latter divergence coincided with species‐level radiations within Homopus (areolatus and femoralis) and Psammobates (oculifer, geometricus and tentorius). Our phylogeny could not resolve relationships within Psammobates, indicating rapid speciation between the Late Oligocene and Early Miocene. The Chersobius species were the last to diverge in the Early to Mid‐Miocene. By the Mid‐Miocene, P. tentorius started to differentiate into four lineages instead of the three recognized subspecies: P. t. tentorius, P. t. trimeni and two P. t. verroxii subclades occurring north and south of the Orange River, respectively. Terminal radiations in several taxa suggest the existence of cryptic species and a more diverse tortoise fauna than currently recognized. Factors contributing to this diversity may include the early origin of African testudinids and climatic fluctuations over a heterogeneous landscape.     

Cover Caption

Cockroaches were dominant insects in diverse Palaeozoic and Mesozoic localities. After the boundary with the Cenozoic (when dinosaurs went extinct), cockroaches are nearly absent and the first significant locality is the Green River in Colorado, USA. It indicates the Eocene cockroaches belong to living genera. The same cosmopolitan fauna is characteristic for the Mexican amber, but after that (possible due to Mid‐Miocene disruption) it was replaced by contemporary cockroach fauna in both Americas. See pages 143–152. Photo provided by Peter Vr?anský     

Geographic and stratigraphic distributions of the Caribbean species of Cladocora (Scleractinia, Faviidae)

A complete account of the faviid genus Cladocora within the Caribbean is presented. In the Caribbean this genus represents an extant group that had its earliest occurrence during the Campanian-Maastrichtian of Jamaica. Recent forms have been reported throughout the Caribbean. The following forms were found (with stratigraphic ranges in the Caribbean): C. arbuscula (Pliocene-Recent), C. debilis (Pleistocene-Recent), C. gracilis (Middle-Upper Maastrichtian), C. jamaicaensis (Campanian-Maastrichtian and Eocene), C. johnsoni (Pliocene), and C. recrescens (Middle-Upper Oligocene). The occurrence of the genus Cladocora in the Caribbean is largely continuous from the Campanian to Recent, during the majority of the Caribbean species show affinities to European assemblages. For the time intervals Paleocene, Lower Oligocene, and Miocene the taxon has not been reported from the Caribbean.     

Integrating fossils in a molecular‐based phylogeny and testing them as calibration points for divergence time estimates in Menispermaceae

Abstract The phylogeny of extant Menispermaceae (Ranunculales) is reconstructed based on DNA sequences of two chloroplast genes (rbcL and atpB) from 94 species belonging to 56 genera. Fossilized endocarps represent 34 genera. The positions of these are inferred using 30 morphological characters and the molecular phylogeny as a backbone constraint. Nine of the thirteen nodes that are each dated by a fossil are used as calibration points for the estimates of molecular divergence times. BEAST is used to estimate stem age (121.2 Myr) and crown age (105.4 Myr) for Menispermaceae. This method does not require an input tree topology and can also account for rate heterogeneity among lineages. The sensitivity of these estimates to fossil constraints is then evaluated by a cross‐validation procedure. The estimated origin for Menispermaceae is dated to the mid‐Jurassic if the customary maximum age of 125 Myr for eudicots is not implemented. All constraints when used alone failed to estimate node ages in some parts of the tree. Fossils from the Palaeocene and Eocene impose strict constraints. Likewise, the use of Prototinomiscium as a dating constraint for Menispermaceae appears to be a conservative approach.