Molecular phylogenetic analyses of shallow-water Caribbean octocorals

Octocorals, especially gorgonians, are conspicuous on Caribbean coral reefs, but there is no consensus regarding species relationships. Mitochondrial protein-coding genes [NADH-dehydrogenase subunits 2 (ND2) and 6 (ND6), and mutS homolog (msh1), 1633 bp] from 28 shallow-water species were sequenced to develop the first molecular phylogeny for Caribbean octocorals. The specimens were collected primarily in the Caribbean or off Brazil in 1999-2001. Morphological characters (sclerites and axial ultrastructure) were also examined in order to map them onto the molecular phylogeny. Analyses of both nucleotide and amino acid substitutions using maximum parsimony and likelihood (including maximum-likelihood and Bayesian analysis) generated very similar results, with most nodes having high levels of support. These molecular results were significantly different from the generally accepted classification. Neither Plexauridae nor Gorgoniidae were monophyletic. Plexaurella spp., nominal plexaurids, were basal to the gorgoniids, sharing many morphological characters with them. This corroborates previous findings using secondary metabolites and biosynthetic pathways. The sea fans, Gorgonia spp. and Pacifigorgia spp., as well as the pinnate gorgonians, Muriceopsis flavida and Pseudopterogorgia spp., did not have sea fan or pinnate relatives, suggesting there has been convergent evolution of colony form. Caribbean plexaurids appeared more derived and/or recently evolved according to both morphological and molecular data (e.g. Eunicea spp. and Plexaura spp.). Molecular phylogenetics is a promising approach for reconstructing phylogenetic relationships among octocorals as well as to understand their complex morphology. Electronic Supplementary Material is available if you access this article at http://dx.doi.org/10.1007/s00227-003-1018-7. On that page (frame on the left side), a link takes you directly to the supplementary material.     

Malaria infection and host behavior: a comparative study of Neotropical primates

Parasites are ubiquitous in populations of free-ranging animals and impact host fitness, but virtually nothing is known about the factors that influence patterns of disease risk across species and the effectiveness of behavioral defenses to reduce this risk. We investigated the correlates of malaria infection (prevalence) in Neotropical primates using data from the literature, focusing on host traits involving group size, body mass, and sleeping behavior. Malaria is spread to these monkeys through anopheline mosquitoes that search for hosts at night using olfactory cues. In comparative tests that used two different phylogenetic trees, we confirmed that malaria prevalence increases with group size in Neotropical primates, as suggested by a previous non-phylogenetic analysis. These results are consistent with the hypothesis that larger groups experience increased risk of attack by mosquitoes, and counter to the hypothesis that primates benefit from the encounter-dilution effect of avoiding actively-seeking insects by living in larger groups. In contrast to non-phylogenetic tests, body mass was significant in fewer phylogeny-based analyses, and primarily when group size was included as a covariate. We also found statistical support for the hypothesis that sleeping in closed microhabitats, such as tree holes or tangles of vegetation, reduces the risk of malaria infection by containing the host cues used by mosquitoes to locate hosts. Due to the small number of evolutionary transitions in sleeping behavior in this group of primates, however, this result is considered preliminary until repeated with a larger sample size. In summary, risk of infection with malaria and other vector-borne diseases are likely to act as a cost of living in groups, rather than a benefit, and sleeping site selection may provide benefits by reducing rates of attack by malaria vectors.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Crinoid phylogeny: a preliminary analysis (Echinodermata: Crinoidea)

We describe the first molecular and morphological analysis of extant crinoid high-level inter-relationships. Nuclear and mitochondrial gene sequences and a cladistically coded matrix of 30 morphological characters are presented, and analysed by phylogenetic methods. The molecular data were compiled from concatenated nuclear-encoded 18S rDNA, internal transcribed spacer 1, 5.8S rDNA, and internal transcribed spacer 2, together with part of mitochondrial 16S rDNA, and comprised 3,593 sites, of which 313 were parsimony-informative. The molecular and morphological analyses include data from the bourgueticrinid Bathycrinus; the antedonid comatulids Dorometra and Florometra; the cyrtocrinids Cyathidium, Gymnocrinus, and Holopus; the isocrinids Endoxocrinus, and two species of Metacrinus; as well as from Guillecrinus and Caledonicrinus, whose ordinal relationships are uncertain, together with morphological data from Proisocrinus. Because the molecular data include indel-rich regions, special attention was given to alignment procedure, and it was found that relatively low, gene-specific, gap penalties gave alignments from which congruent phylogenetic information was obtained from both well-aligned, indel-poor and potentially misaligned, indel-rich regions. The different sequence data partitions also gave essentially congruent results. The overall direction of evolution in the gene trees remains uncertain: an asteroid outgroup places the root on the branch adjacent to the slowly evolving isocrinids (consistent with palaeontological order of first appearances), but maximum likelihood analysis with a molecular clock places it elsewhere. Despite lineage-specific rate differences, the clock model was not excluded by a likelihood ratio test. Morphological analyses were unrooted. All analyses identified three clades, two of them generally well-supported. One well-supported clade (BCG) unites Bathycrinus and Guillecrinus with the representative (chimaeric) comatulid in a derived position, suggesting that comatulids originated from a sessile, stalked ancestor. In this connection it is noted that because the comatulid centrodorsal ossicle originates ontogenetically from the column, it is not strictly correct to describe comatulids as unstalked crinoids. A second, uniformly well-supported clade contains members of the Isocrinida, while the third clade contains Gymnocrinus, a well-established member of the Cyrtocrinida, together with the problematic taxon Caledonicrinus, currently classified as a bourgueticrinid. Another cyrtocrinid, Holopus, joins this clade with only weak molecular, but strong morphological support. In one morphological analysis Proisocrinus is weakly attached to the isocrinid clade. Only an unusual, divergent 18S rDNA sequence was obtained from the morphologically strange cyrtocrinid Cyathidium. Although not analysed in detail, features of this sequence suggested that it may be a PCR artefact, so that the apparently basal position of this taxon requires confirmation. If not an artefact, Cyathidium either diverged from the crinoid stem much earlier than has been recognised hitherto (i.e., it may be a Palaeozoic relic), or it has an atypically high rate of molecular evolution.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Thorpe, Port Erin     

Molecular evidence for multiple lineages in the gorgonian family Plexauridae (Anthozoa: Octocorallia)

Octocorals are diverse and abundant on many marine hard substrates, and, within this group, members of the family Plexauridae are an important component of tropical reef assemblages, especially in the Caribbean. To understand historical relationships within this large and diverse assemblage, and to test the monophyly of the family and some of its genera, DNA sequences of two mitochondrial loci (msh1 and ND2, ~1,185 bp) were analyzed from 46 species in 21 genera from deep and shallow waters in the tropical western Atlantic and in the tropical western and eastern Pacific (plus 9 taxa in the closely related Gorgoniidae and 1 species of the more distantly related Alcyoniidae). Five strongly supported clades were recovered. Three large clades correspond roughly to the Plexauridae, Paramuriceidae, and Gorgoniidae, and two smaller clades were comprised of taxa previously assigned to several families. Astrogorgia sp. did not group with any of the clades. The mutual relationships among the five clades remain unclear. Several genera previously regarded as unrelated appear to be grouped among the three families; e.g. Hypnogorgia sp. (Paramuriceidae) falls within a clade consisting of both Pacific and Atlantic Muricea spp. (Plexauridae), while Swiftia sp., Scleracis sp., and an Atlantic Thesea sp. (all Paramuriceidae) group with the gorgoniids. In several instances, genera containing Atlantic and Pacific species were recovered as monophyletic (Muricea spp., Bebryce spp.). However, in at least three cases (Echinomuricea spp., Thesea spp., Villogorgia spp.), placement of Atlantic and Pacific species in the same genus may reflect convergence of sclerite morphology. The results indicate a strong need for reexamination of octocoral taxonomy using a combination of molecular, morphological, and chemical evidence.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-005-1592-y.Communicated by J.P. Grassle, New Brunswick     

Anatomical and experimental evidence for particulate feeding in<Emphasis Type="Italic"> Lucinoma aequizonata</Emphasis> and<Emphasis Type="Italic"> Parvilucina tenuisculpta</Emphasis> (Bivalvia: Lucinidae) from the Santa Barbara Basin

Previous nutritional models for adults of the lucinid bivalve Lucinoma aequizonata contend that symbiotic chemoautotrophic bacteria provide most of the organic carbon for the host. The existence of this symbiosis, coupled with the hosts distinctive anatomical features, shaped the impression that particulate feeding was not a significant part of L. aequizonata nutrition. Here, we use several techniques to show that particulate feeding is a consistent and important part of the L. aequizonata nutritional strategy. Histological and scanning electron microscopy observations reveal that the gills of L. aequizonata, like those of the lucinid Parvilucina tenuisculpta, have functional mucociliary epithelia, able to transport captured particles to the mouth. Observations of gut content and radiolabeled feeding experiments indicate that L. aequizonata does ingest and assimilate carbon from particulate organic matter. Furthermore, molecular identification of a broad spectrum of organisms in the guts of native adult specimens demonstrates that L. aequizonata is non-selective when ingesting organic material, and has a mixotrophic diet.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by P.W. Sammarco, Chauvin     

Genetic Criteria for Establishing Evolutionarily Significant Units in Cryan's Buckmoth

Buckmoths ( Hemileuca spp.) are day-flying saturniid moths with diverse ecologies and host plants. Populations that feed on Menyanthes trifoliata , known commonly as Cryan's buckmoths, have been found in only a few bogs and fens near eastern Lake Ontario in New York and near Ottawa in Ontario, Canada. Because of their unique ecological traits, geographic isolation from other Hemileuca populations, and the small number of sites they occupy, there is concern that the Cryan's buckmoth populations are phylogenetically distinct and should be protected. The Cryan's buckmoths have not yet been taxonomically described and do not appear to have clear distinguishing morphological characters. Both molecular genetic traits (allozymes and mitochondrial DNA sequences) and an ecologically based character (host performance) were investigated to determine whether these populations possess fixed diagnostic characters signifying genetic differentiation from other eastern Hemileuca populations. Such differences would merit separate conservation management as an evolutionarily significant unit. Our studies showed that the Cryan's buckmoths clearly belong to the Hemileuca maia species group, but they could not be readily distinguished from other members of that group by means of molecular genetic techniques. There were no fixed differences in alleles or haplotypes distinguishing any of the populations or species, suggesting recent divergence. Nonetheless, in the host-plant performance experiment only the Cryan's buckmoth larvae were able to develop on M. trifoliata , a significant difference from other Hemileuca larvae tested. The Cryan's buckmoth appears to be unique in host performance and warrants protection and management us an evolutionarily significant unit. In cases such as this where groups appear to have recently diverged, investigations into ecologically significant traits may provide indicators of conservation significance as reliable as molecular genetic markers.     

Assessing feeding of a carnivorous copepod using species-specific PCR

The polymerase chain reaction (PCR) offers a sensitive and selective way to detect trace amounts of biological remnants. Here, we show that this simple molecular technique can be applied to identify prey copepods in the fecal pellets of carnivorous zooplankton. Using variation in the mitochondrial cytochrome C oxidase subunit I (mtCOI) sequence, we developed a species-specific oligonucleotide PCR primer (COI-2026) for Calanus helgolandicus. In a touch-down PCR, Calanus DNA was amplified from pellets collected from freshly incubated individuals of the carnivorous copepod Pareuchaeta norvegica. Positive results could easily be detected by agarose gel electrophoresis.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by M. Kühl, Helsingør     

Spatial and temporal variation in the diet of a high trophic level predator,the Australian fur seal (<Emphasis Type="Italic">Arctocephalus pusillus doriferus</Emphasis>)

This study quantifies the manner in which Australian fur seals, Arctocephalus pusillus doriferus, use their prey in a spatial and temporal context. We analysed 977 scat and 66 regurgitate samples collected from Tasmanian breeding colonies and haul-outs between 1994 and 2000. Diagnostic prey remains identified in the scats represented 35 fish taxa and 8 cephalopod taxa. The main taxa identified in scats, where frequency of occurrence was 10%, were leatherjacket species (family Monocanthidae), redbait (Emmelichthys nitidus), barracouta (Thyrsites atun), jack mackerel (Trachurus declivis) and red cod (Pseudophysis bachus). Regurgitates were dominated by cephalopods, primarily Goulds squid (Nototodarus gouldi), Octopus maorum, O. berrima/pallidus and Sepia apama. Discriminant function analyses indicated that there were generally no significant differences in the composition of the diet between colonies within a year, suggesting that prey distribution is fairly uniform throughout Bass Strait at those time scales. The diet at breeding colonies, however, exhibited significant inter- and intra-annual variation, determined by the presence of several key taxa, such as barracouta and a species of scorpionfish (family Scorpaenidae). The diet composition also varied regionally, between Bass Strait and southern Tasmania in spring 1999 and autumn 2000, with redbait, barracouta and a species of scorpionfish identified as the main taxa contributing to this difference. Redbait occurred in the diet only in southern Tasmania, whereas barracouta and scorpionfish occurred only in Bass Strait.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-003-1219-0.Communicated by G.F. Humphrey, Sydney     

Differences in chloroplast ultrastructure of Phaeocystis antarctica in low and high light

Phaeocystis antarctica Karsten exhibits optical changes in pigment packaging during acclimation to drastically different light levels. Here, the three-dimensional morphological rearrangements are shown for two light conditions mimicking limiting and saturating conditions for photosynthesis. Cultures of P. antarctica were grown semi-continuously under light-limiting conditions for growth (14 μmol quanta m⁻² s⁻¹) and a light-saturating condition (259 μmol quanta m⁻² s⁻¹) for growth. Increased numbers of thylakoids were observed under the low light treatment. In contrast, there were less amounts of thylakoid stacking in each chloroplast under the high light treatment. Electron microscopic tomographic reconstructions illustrate the complexity of the chloroplast organelle where the thylakoids ‘interact’ with the pyrenoid and the chloroplast membrane. Highly complex characteristics, such as bi- and tri-furcations in the thylakoid stacks, were continuous throughout the chloroplast. Other organelles, such as the Golgi apparatus and dense vesicles that may potentially affect cellular scattering and absorption were also observed in both high and low light. These three dimensional thylakoid arrangements have profound implications for cellular photophysiology. They represent a new view of algal chloroplast structure, and provide a starting point for more accurate optical modeling studies.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

稀有鮈鲫Dmrt基因家族13个成员的克隆与序列分析

已经发现果蝇Doublesex、线虫Mab-3、青DMRT1Y/DMY和人类DMRT1等性别决定与分化基因均含有一个具有DNA结合能力的保守基序--DM结构域,对性别决定和性别分化具有调控功能.利用简并PCR,从稀有鲫基因组DNA中克隆了13个具有不同DM结构域的Dmrt基因家族成员.基于DM保守基序,建立了各物种的进化树.结果表明,稀有鲫基因组存在多个Dmrt基因成员,该基因在脊椎动物和非脊椎动物中具有高度保守性,是一种理想的环境内分泌干扰物研究的分子模型,在分子生态毒理学研究中具有很好的应用前景.     

Mating behavior of female rock shrimp Rhynchocinetes typus (Decapoda: Caridea)—indication for convenience polyandry and cryptic female choice

While studies on a wide diversity of organisms have demonstrated the importance of female behavior during matings, in crustacean studies, a strong bias towards male mating behavior prevails. Reproductively mature rock shrimp (Rhynchocinetes typus) exist as several ontogenetic stages that differ in their morphological and physiological capacities. In natural populations, the majority of males are in early ontogenetic stages (termed typus), many are in intermediate stages (intermedius), and few are in the terminal molt stage (robustus). Dominant robustus males, which have already demonstrated their biological fitness by surviving to this stage, have previously been shown to have a higher potential than subordinate typus males to defend receptive females against other males, and fertilize the entire clutch of a female. While females should thus show a preference for robustus males, they nevertheless frequently receive sperm from typus males. These observations suggested that females might have mechanisms to discriminate against sperm from subordinate males. In laboratory experiments, we observed that females avoided being seized by typus males for longer time periods in the absence of robustus males than in their presence. Following seizure, females that were initially held by typus males, required more time to initiate spawning than those held by robustus males. Many typus males transferred spermatophores to females before these started to spawn while robustus males waited until females began to spawn before they transferred spermatophores. Females manipulated spermatophores received from typus males for long time periods (minutes), but not those they received from robustus males. By accepting sperm from subordinate typus males, females may avoid further harassment (convenience polyandry), but they subsequently may discriminate against these subordinate males by delaying spawning and removing their sperm. These observations suggest that female behavior influences the outcome of matings, favoring fertilization of eggs by sperm from dominant males. Convenience polyandry and cryptic female choice may be common in other crustaceans as well.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by T. Czeschlik     

Utilization of a new host in the screaming cowbird <Emphasis Type="Italic">Molothrus rufoaxillaris</Emphasis>, a host specialist brood parasite: host switch or host acquisition?

The screaming cowbird Molothrus rufoaxillaris has been long known as a host specialist brood parasite. However, in the past years, the utilization of two new hosts has been documented. We examined the variation in mitochondrial control region sequences from screaming cowbird chicks found in the nests of two hosts, the bay-winged cowbird (Agelaioides badius), which is its regular host, and the chopi blackbird (Gnorimopsar chopi), which is a new host, in Formosa Province, Argentina. If a group of females switched to this new host, we expected to find an association between host use and haplotype frequency distribution, indicating the presence of host-specific female lineages, whereas we expected no such association if the cowbird population incorporated this new host and females use both hosts simultaneously. Haplotype frequency distributions differed between cowbird chicks from the nests of both hosts. This indicates that nest choice by females of this brood parasite is not random and that they preferentially parasitize the nests of the same host species.     

Survey of the occurrence and morphology of sub-cuticular bacteria in shelf echinoderms from the north-east Atlantic Ocean

Echinoderms from the shelf seas around the British Isles were examined by epifluorescence and transmission electron microscopy for the presence of sub-cuticular bacteria (SCB). Information was obtained on SCB distribution in 63 of the 88 species known from this area, and 40 of these contained SCB. The SCB were present in all five classes. Only 39% of the holothurian species had SCB. In each of the other four classes, 60% of the species were associated with SCB. No correlation was noted between host ecology and SCB distribution or morphology. SCB distribution appeared to be related to host phylogeny. Congeneric echinoderms almost always had or lacked SCB, and this was usually also true for co-familial species. SCB showed limited morphological diversity, with all of them fitting into a classification of three general types. There was no morphological evidence suggesting that they are chemoautotrophs or methylotrophs. Some species had two morphological types of symbiont. SCB load was quantified for four species, three of which had symbiont densities >10⁹ SCB g^-1 ash-free dry wt. SCB are probably heterotrophic and may be co-evolving with their hosts.     

Two-dimensional wavelet analysis of spruce budworm host basal area in the Border Lakes landscape

Increases in the extent and severity of spruce budworm (Choristoneura fumiferana Clem.) outbreaks over the last century are thought to be the result of changes in forest structure due to forest management. A corollary of this hypothesis is that manipulations of forest structure and composition can be used to reduce future forest vulnerability. However, to what extent historical forest management has influenced current spatial patterns of spruce budworm host species is unknown. To identify landscape-scale spatial legacies of forest management in patterns of spruce budworm host species (i.e., Abies balsamea and Picea spp.), we analyzed remotely sensed forest data from the Border Lakes landscape of northern Minnesota and northwestern Ontario. Our study area contains three regions with different management histories: (1) fine-scale logging patterns in Minnesota, (2) coarse-scale logging patterns in Ontario, and (3) very limited logging history in the Boundary Waters Canoe Area and adjacent Quetico Provincial Park. We analyzed forest basal-area data using wavelets and null models to identify: (1) at which scales forest basal area is structured, (2) where those scales of pattern are significantly present, and (3) whether regions of local significance correspond to regional boundaries that separate the study area. Results indicate that spatial patterns in host basal area are created by nonstationary processes and that these processes are further constrained by lakes and wetlands. Wavelet analysis combined with significance testing revealed a bimodal distribution of scale-specific wavelet variance and separate zones of host species basal area that partially correspond with regional boundaries, particularly between Minnesota and the Wilderness region. This research represents one of the first comparisons of forest spatial structure in this region across an international border and presents a novel method of two-dimensional wavelet analysis that can be used to identify significant scale-specific structure in spatial data.     

Molecular characterization of the zoanthid genus <Emphasis Type="Italic">Isaurus</Emphasis> (Anthozoa: Hexacorallia) and associated zooxanthellae (<Emphasis Type="Italic">Symbiodinium</Emphasis> spp.) from Japan

The zoanthid genus Isaurus (Anthozoa: Hexacorallia) is known from both the Indo-Pacific and Atlantic Oceans, but phylogenetic studies examining Isaurus using molecular markers have not yet been conducted. Here, two genes of markers [mitochondrial cytochrome oxidase subunit I (COI) and mitochondrial 16S ribosomal DNA (mt 16S rDNA)] from Isaurus specimens collected from southern Japan (n = 19) and western Australia (n = 3) were sequenced in order to investigate the molecular phylogenetic position of Isaurus within the order Zoantharia and the family Zoanthidae. Additionally, obtained sequences and morphological data (polyp size, mesentery numbers, mesogleal thickness) were utilized to examine Isaurus species diversity and morphological variation. By comparing our obtained sequences with the few previously acquired sequences of genera Isaurus as well as with Zoanthus, Acrozoanthus (both family Zoanthidae), and Palythoa spp. (family Spenophidae) sequences, the phylogenetic position of Isaurus as sister to Zoanthus within the Family Zoanthidae was suggested. Based on genetic data, Isaurus is most closely related to the genus Zoanthus. Despite considerable morphological variation (in particular, polyp length, mesentery numbers, external coloration) between collected Isaurus specimens, all specimens examined are apparently conspecific or very closely related based on molecular data and observed morphological variation within colonies. Additionally, obtained internal transcribed spacer of ribosomal DNA (ITS-rDNA) sequences from symbiotic zooxanthellae (Symbiodinium spp.) from all Isaurus specimens were shown to be subclade C1-related Symbiodinium. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Carotenoid-derived ornaments reflect parental quality in male and female yellow-eyed penguins (<Emphasis Type="Italic">Megadyptes antipodes</Emphasis>)

The handicap principle suggests that ornamental traits that function as honest signals in mate selection must be costly to be effective. We evaluated in the sexually monochromatic yellow-eyed penguin (Megadyptes antipodes) whether the carotenoid-derived plumage and eye coloration predicts parental quality and whether males and females within pairs mate assortatively in relation to these carotenoid-derived ornaments. In addition, we investigated whether age or body condition was related to the coloration of the ornamental traits. In yellow-eyed penguins, parental quality of males and females was predicted by eye and head plumage coloration. Even when we controlled for gender- and age-specific differences, eye and head plumage coloration reflected honestly parental quality. Males and females mated assortatively in relation to these ornamental traits. While age influenced coloration of both the eye and head plumage, body condition was related only to the saturation of plumage coloration. These results provide evidence that the carotenoid-derived ornaments in yellow-eyed penguins reflect the parental abilities of birds and, therefore, may be costly signals. Potentially, female and male yellow-eyed penguins could use eye and plumage coloration as an indirect cue in assessing age and quality of individual birds during mate choice. This is only the second study to examine plumage coloration in relation to sexual selection in penguins, while conspicuous ornamental traits in other species of penguin beg the question whether they also play a role in sexual selection.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by C.R. Brown     

High intervality explained by phylogenetic constraints in host-parasite webs

The finding of invariant structures in species interaction webs is of central importance for ecology, with the greatest challenge remaining the elucidation of the processes governing these universal web patterns. Here we quantify the degree of intervality of seven fish-metazoan and 33 mammal-flea webs, i.e., the number of irreducible gaps in parasite diets along the host spectrum, and then challenge the idea that some invariant structures may emerge in host-parasite webs. Using a null model of random links between parasite and host species we find that empirical host-parasite webs exhibit a strong bias toward contiguity of parasite diet, i.e., toward intervality. Going one step further, we demonstrate that a null model with phylogenetic constraints on host-parasite links produced webs very similar to empirical ones, particularly when phylogenetic constraints occur at the family level, that is, when two hosts from the same family are more likely to be infected than two random hosts. In addition, we propose a new standardized measure of intervality which describes a novel "facet" of natural networks as it is independent of connectance or web size. We suggest using this measure as a surrogate of web maturity or saturation as phylogenetic constraints can drive webs toward intervality.