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Figure 16. A-C in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 16. A-C, Macropora levinseni Brown, Recent, New Zealand, NIWA. A, partly bleached ovicell, viewed laterally. B, interior of brooding cavity, showing bases of ooecial costae overgrown by cryptocyst; gymnocystal ovicell floor is below. C, ovicell upside down, showing the pores in its floor. D-F, Macropora polymorpha (Philipps). Recent, New Zealand, NIWA. D, developing ovicell, showing gymnocystal floor and intercostal spaces with cryptocystal fabric. E, upside-down ovicell, showing the kenozooidal supporting zooid (top) and peripheral pores in the ovicell floor. F, part of the developing ovicell, showing costal coelomic lumina connected with lacunae in the cryptocystal matrix. Scale bars: A, D = 500 Mm; B, F = 100 Mm; C, E = 200 Mm.
Figure 2. Heteroecium amplectens Hincks, NHM 99.5.1.702 in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 2. Heteroecium amplectens Hincks, NHM 99.5.1.702, Recent, Western Australia. A, part of a colony with several nonbrooding zooids and one brooding zooidal complex. B, brooding zooidal complex. C, brooding zooidal complex from below (distal to the right); openings of the costae surround the floor of brood chamber. D, membranous area with two appendages in the brood-chamber floor; a communication pore in the transverse wall between the maternal zooid and distal kenozooiod can be seen in the left lower corner. Scale bars: A, C = 100 Mm; B, D = 50 Mm.
Figure 11. A-C in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 11. A-C, Monoporella sp. 1, Recent, Japan, Pacific Ocean, GSUH. A, oblique lateral view of bleached colony fragment with an ovicell; note distal fissures and large foramina. B, internal ooecial wall, showing fused costae and slits between their bases covered with cryptocyst. C, broken costal bases embedded in cryptocyst. D-F, Monoporella nodulifera (Hincks), Recent, Australia; unbleached broken ovicell roof upside down; membranous walls are seen on both sides, as well as three flattened costae covered by cryptocyst which is pierced by canals; cryptocystal insertions with pores are seen between the costae. Scale bars: A = 200 Mm; B, D, E, F = 50 Mm; C = 10 Mm.
Figure 5. A, B in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 5. A, B, Unidistelopora krauseae (Voigt & Schneemilch), Lower Campanian, northern Germany, VC T10580. A, maternal zooid with ovicell and intramural bud. B, ovicell spine bases arranged in a semicircle along a ridge on the proximal gymnocyst of the distal zooid. C-F, Gilbertopora larwoodi Ostrovsky & Taylor, Lower Cenomanian, Cambridge, England. C, complete ovicell of two flattened spines; NHM D23297. D, complete ovicell viewed from the side, showing a lateral foramen; NHM D23298. E, complete ovicell in proximal view, showing the main opening of the ovicell; NHM D23298. F, complete ovicell in distal view, showing the distal opening; NHM D23298. Scale bars: A = 100 Mm; B-F = 50 Mm.
Figure 23 in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 23. Schematic diagrams of brood chambers in Monoporellidae (A, C, D) and Macroporidae (B, E) in longitudinal and transverse section, showing maternal and distal zooids; cryptocystal ooecial matrix is shadowed. A, Monoporella sp. 1. B, Macropora sp. 1 and M. cribrilifera (transverse section made through cryptocystal (left) and gymnocystal (right) components of the ribs). C, Monoporella sp. 2. D, Monoporella nodulifera. E, Macropora levinseni.
Figure 15. A, B in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 15. A, B, Macropora sp. 1., Lower Miocene, Auckland, New Zealand. A, colony with one complete and one damaged ovicell; NHM BZ5202. B, broken ovicell; NHM BZ5203. C, Macropora sp. 2, Recent, Cavalli Seamounts, New Zealand, NIWA. Ovicell from the inside, showing gymnocystal costal surfaces and intercostal spaces with cryptocystal fabric and fissures between them. D-F, Macropora uttleyi López de la Cuadra & García Gómez, Recent, Cavalli Seamounts, Pacific Ocean, NIWA. D, ovicellate zooid. E, broken ovicell, showing gymnocystal floor and inner costal surfaces. F, detail of ovicell interior, showing flat-surfaced costae and intercostal spaces with pores and cryptocystal fabric. Scale bars: A = 500 Mm; B, C = 100 Mm; D, E = 200 Mm; F = 50 Mm.
Figure 10. A, B in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 10. A, B, Monoporella multilamellosa (Canu & Bassler), Eocene, North Carolina, USA, NHM BZ4860. A, complete ovicell consisting of two flattened and expanded spines overgrown distally by a narrow fringe of cryptocyst; note lateral foramina. B, broken ovicell exposing gymnocystal floor and showing the medial gap between the bases of the two ovicell spines. C-F, Monoporella sp. 2, Recent, Alaska. C, bleached colony fragment with two complete ovicells and lateral foramen arrowed; MNHN 2856–7(b). D, unbleached ovicell; MNHN 2856–7(a). E, broken costa showing coelomic lumen; MNHN 2856–7(b). F, oblique view showing gymnocystal internal surface of ovicell; note limit of the cryptocystal expansion from the outer ovicell surface, and longitudinal grooves; MNHN 2856–7(b). Scale bars: A, B, E, F = 100 Mm; C = 500 Mm; D = 200 Mm.
Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489 in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm.
Figure 22 in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 22. Schematic diagrams of brood chambers in Monoporellidae (A-C), Cribrilinidae (D, E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Stichomicropora spp. with articulated ovicell spine bases; B, Stichimicropora baccata. C, Monoporella multilamellosa. D, Leptocheilopora spp. E, Bellulopora bellula.
Figure 3 in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 3. Distelopora bipilata Lang, Lower Cenomanian, Cambridge, England. A, part of a colony with several nonovicellate autozooids and one broken ovicell, NHM D21883. B, ovicell spine bases forming a semicircle, NHM D21881. C, ovicell spine bases at some distance from the mural rim, NHM D21881. D, ovicell spine bases forming a gentle arch. Medial spines are adjacent to the proximal edge of the mural rim of the distal zooid, NHM BZ4958. Scale bars: A = 100 Mm; B–D = 50 Mm.
Figure 14. A–C in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 14. A–C, Macropora cribrilifera Maplestone, Lower Miocene, South Australia, NMV P311815. A, ovicell in oblique frontal view. B, ovicell viewed from the distal side (orifice of distal zooid bottom centre). C, lateral view of ovicell, showing intercostal slits and pores. D-F, Macropora waimatukuensis (Uttley). CM zb51,?Miocene, Southland, New Zealand. D, complete ovicell; note calcified opercula in distal and two lateral zooids. E, complete and broken ovicells. F, proximal part of ovicell showing cryptocystal costal fabric. Scale bars: A-D = 200 Mm; E = 500 Mm; F = 100 Mm.
Figure 21 in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi.
Figure 7. A, B in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 7. A, B, Stichomicropora marginula (Brydone), Coniacian, Kent, England, NHM D44609. A, part of colony with ovicellate and nonovicellate zooids. B, maternal zooid with ovicell preserved as a gently curved, distally convex arch of spine bases. C, D, Stichomicropora sp. 1, Campanian, Norwich, England, NHM D42263. C, several fertile zooids with ovicells represented by spine bases arranged in distally concave or distally convex gentle arches, or in a straight line. D, view centred on a damaged part of a colony with distally convex rows of ovicell spine bases (upper left and upper right) and a distally concave row (bottom right). E, F, Stichomicropora sp. 2, Campanian, Clarendon, England, NHM D46004. E, edge of colony, showing three ovicells, represented by gently curved, distally convex arches of spine bases, in zooids with broken frontal shields. F, ovicell spine bases and floor. Scale bars: A = 500 Mm; B, F = 100 Mm; C, E = 250 Mm; D = 200 Mm.
Figure 17 in Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans
Figure 17. Macropora levinseni Brown, Recent, New Zealand. A, longitudinal section of an ovicell, showing costal lumen (arrowed) and zooidal operculum (right of arrow). B, saggital section of ovicell with embryo, showing attachments (arrowed) of the internal membranous ooecial wall to the calcified part of the ooecium; note thick external membranous ooecial wall. Scale bars = 100 Mm.
Fig. 6 in Brooding behaviour of the centipede Otostigmus spinosus Porat, 1876 (Chilopoda: Scolopendromorpha: Scolopendridae) and its morphological variability in Thailand
Fig. 6. Variability in taxonomic characters in Otostigmus spinosus Porat, 1876 (examination materials: CUMZ 00224, Wat Tham Wararam, Phanom district, Surat Thani; CUMZ 00229, Tham Wang Thong, Phatthalung; CUMZ 00231, Surin Islands, Phang-nga). A, Ventral view of cephalic plate and antenna; B, Dorsal view of cephalic plate and antenna; C, Coxosternite; D–F, Variation in coxosternal teeth; G, Surface of TT5–7; H, Variability of depressions on surface of sternites 5–7; I, Leg 20 with projection on distal end of tarsus 1; J, Leg 21 with projection on tarsus 1; K, Ultimate leg-bearing segment, ventral view, with arrangement of ventral spines on prefemur of ultimate legs; L, Tergite of ultimate leg-bearing segment with arrangement of dorsal and corner spines on prefemur of ultimate legs; M, Pore field on coxopleuron (left).
Fig. 5. Otostigmus spinosus Porat, 1876 in Brooding behaviour of the centipede Otostigmus spinosus Porat, 1876 (Chilopoda: Scolopendromorpha: Scolopendridae) and its morphological variability in Thailand
Fig. 5. Otostigmus spinosus Porat, 1876. Light photographs of specimens from southern Thailand (Fig. A–E; specimens from Surin Islands, Phang-nga province: CUMZ 00231; Fig. I–N; specimens from Wat Tham Wararam, Surat Thani province: CUMZ 00224). A, Cephalic plate and T1 with first pair of locomotory legs; B, Forcipular segment; C, Antenna with cephalic plate and TT1–5; D, Tergites 9–10; E, Sternites 9–10; F–H. Spiracles on segments 3, 5 and 8, respectively; I, Tergite and sternite of ultimate-leg bearing segment; J, Pore field on left and right coxopleura; K, Dorsal view of ultimate leg prefemur; L, Ventral view of ultimate leg prefemur; M–N, Leg 20 left and right with spine on distal part of prefemora.
Data for: Brood parasitism of Hooded Warblers by Brown-headed Cowbirds: Severe impact on individual nests but modest consequences for seasonal fecundity and conservation
<div> <div> <div> <div> <p>Brood parasitism by Brown-headed Cowbirds (<em>Molothrus ater</em>) often has pronounced negative effects on host nests. However, the extent to which parasitism reduces annual reproduction and presents conservation challenges for host species is unclear. We address this issue with data from a color-banded population of Hooded Warblers (<em>Setophaga citrina</em>) in Pennsylvania, where Hooded Warblers have increased dramatically despite frequent nest parasitism. Our analysis is based on both an extensive dataset (8 years, 847 nests) on the per-nest impacts of cowbird parasitism, and female-based stochastic simulations that accurately reflect the reproductive biology and parasitism rate (30%) of our study population. Cowbird parasitism has multiple negative consequences for Hooded Warbler nests, including: (1) reduced host clutch size, (2) increased nest abandonment, (3) increased risk of complete failure due to predation, and (4) in surviving nests increased egg loss, hatching failure, and nestling mortality. We estimate that parasitism reduces success of Hooded Warbler nests 68%, from 1.29 to 0.41 fledglings per nest. For females and populations, however, the consequences of nest parasitism are considerably less extreme; female annual fecundity decreases 25% for each nesting attempt parasitized, and population-level fecundity drops 5.6% for each 10% increase in the frequency of parasitism. These more modest impacts are attributable to: (1) steep declines in rates of cowbird parasitism as the nesting season progresses, (2) rapid re-nesting following abandonment or failure of parasitized nests, and (3) regular double brooding, with second broods initiated in late June and July when the incidence of cowbird parasitism is low. Our results help resolve the paradox of how cowbird parasitism can have both severe consequences for individual host nests but more modest and sustainable conservation impacts on the seasonal fecundity of females and populations. They further underscore the importance of determining population-level effects of brood parasites before investing in costly management efforts.</p> </div> </div> </div> </div>
Rapid evolution of a brood parasite's egg pattern does not lead to large increases in mimetic fidelity
<p>In coevolutionary mimicry systems, mimics evolve to resemble models. When model fitness is reduced by close mimicry (such as when models are hosts and mimics are brood parasites), selection should drive mimics to evolve greater similarity to models over time, and models to evolve away from mimics ('chase-away evolution'), potentially resulting in mimetic fidelity remaining constant over time. Evidence for reciprocal evolution of models and mimics and its effect on mimetic fidelity is limited, however, likely because long-term data are required to observe such trends. Here, we test for these dynamics in an avian brood parasite system. The brood-parasitic cuckoo finch (Anomalospiza imberbis) lays eggs which mimic those of its host, the tawny-flanked prinia (Prinia subflava). In defence, prinias have evolved complex egg patterns which facilitate recognition of parasitic eggs. Prinia egg patterns are more complex than cuckoo finch egg patterns, and so selection should favour elevated complexity in parasite eggs (i.e. mimics evolving towards models) and even greater complexity in host eggs (i.e. models evolving away from mimics). Using a dataset spanning 50 years, we show that egg pattern complexity has indeed increased in both species over this time period, reflecting rapid adaptive evolution of both models and mimics. Both species have evolved at similar rates. Accordingly, we find no detectible increase in mimetic fidelity over time. Thus, host evolution can counteract even rapid parasite evolution, and result in the persistence of imperfect mimicry.</p>
Combined measures of mimetic fidelity explain imperfect mimicry in a brood parasite–host system
<p>The persistence of imperfect mimicry in nature presents a challenge to mimicry theory. Some hypotheses for the existence of imperfect mimicry make differing predictions depending on how mimetic fidelity is measured. Here, we measure mimetic fidelity in a brood parasite–host system using both trait-based and response-based measures of mimetic fidelity. Cuckoo finches <em>Anomalospiza</em> <em>imberbis</em> lay imperfectly mimetic eggs that lack the fine scribbling characteristic of eggs of the tawny-flanked prinia <em>Prinia</em> <em>subflava</em>, a common host species. A trait-based discriminant analysis based on Minkowski functionals—that use geometric and topological morphometric methods related to egg pattern shape and coverage—reflects this consistent difference between host and parasite eggs. These methods could be applied to quantify other phenotypes including stripes and waved patterns. Furthermore, by painting scribbles onto cuckoo finch eggs and testing their rate of rejection compared to control eggs (i.e. a response-based approach to quantify mimetic fidelity), we show that prinias do not discriminate between eggs based on the absence of scribbles. Overall, our results support relaxed selection on cuckoo finches to mimic scribbles, since prinias do not respond differently to eggs with and without scribbles, despite the existence of this consistent trait difference.</p>
Low incidence of sibling cannibalism among brood parasitic cuckoo catfish embryos
<p>Brood parasites have demanding needs of host resources. Brood parasitic offspring are highly competitive and frequently cause the failure of host broods and the survival of a single parasitic offspring. Accordingly, virulent brood parasites lay single eggs in host nests to minimize multiple parasitism and ensuing sibling competition. In the cuckoo catfish (<em>Synodontis multipunctatus</em>), which parasitise mouthbrooding cichlid fishes in Lake Tanganyika, the modes of host and parasite oviposition lead to frequent cases of multiple parasitism. We experimentally tested the prediction that multiple parasitism leads to frequent siblicide. Cuckoo catfish embryos prey upon host offspring to obtain nourishment during their 3-week development in the host buccal cavity and may also consume conspecific siblings. The potential benefits of siblicide in the system are, therefore, twofold: to decrease competition for limited resources (i.e. host brood with rich yolk sacs) and to directly obtain nourishment by consuming rivals. We found that sibling cannibalism indeed provided measurable benefits in terms of increased growth of the cannibals, but sibling cannibalism was rare and typically occurred only when all host offspring had been consumed. This implies that cannibalism in the cuckoo catfish embryos emerges to mitigate starvation rather than eliminate sibling competition. </p>
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